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The feeding biology of tintinnid protozoa and some other inshore microzooplankton Blackbourn, David John 1974

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THE FEEDING BIOLOGY OF TINTINNID PROTOZOA AND SOME OTHER INSHORE MICROZOOPLANKTON by David John Blackbourn A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY i n the Department of Zoology and I n s t i t u t e of Oceanography We accept t h i s t h e s i s as conforming to the required standard THE UNIVERSITY OF BRITISH COLUMBIA September, 1974 > In presenting th i s thes is in pa r t i a l fu l f i lment of the requirements for an advanced degree at the Univers i ty of B r i t i s h Columbia, I agree that the L ibrary sha l l make it f ree ly ava i lab le for reference and study. I fur ther agree that permission for extensive copying of th i s thes is for scho lar ly purposes may be granted by the Head of my Department or by his representat ives. It is understood that copying or pub l i ca t ion of th is thes is for f inanc ia l gain sha l l not be allowed without my writ ten permission. Department of ^ O o C o G Y The Univers i ty of B r i t i s h Columbia Vancouver 8, Canada . Date i ABSTRACT Tin t i n n i d s are among the largest and most abundant of the marine c i l l a t e microzooplankton but there i s very l i t t l e published information on t h e i r feeding rates and a b i l i t i e s . The feeding of Tintinnopsis subacuta (and to a l e s s e r extent, that of 12 other species) was investigated with three methods 1) d i r e c t observation 2) counts of accumulated food c e l l s and 3) Coulter Counts of the p a r t i c l e s i n the experimental medium. There was reasonable q u a l i -t a t i v e agreement between the r e s u l t s obtained by the three methods but quantitative agreement was poor. Many of the r e s u l t s showed no s i g n i -f i c a n t differences due to very great v a r i a b i l i t y i n the r e s u l t s for a singl e t i n t i n n i d species within and between experiments. Much of t h i s v a r i a b i l i t y may be due to the methods used but i t also r e f l e c t s the v a r i -a b i l i t y of t i n t i n n i d s i n natural populations. A wide v a r i e t y of items was eaten by t i n t i n n i d s , including smaller t i n t i n n i d s ; and the maximum food s i z e can be rela t e d to t i n t i n n i d c e l l volume over a wide range but i s d i s s i m i l a r i n t i n t i n n i d species of s i m i l a r c e l l s i z e . . Several t i n t i n n i d species showed d i f f e r e n t i a l pre-dation on various types of laboratory phytoplankton. This d i f f e r e n t i a l predation was based upon the a b i l i t y of the predator to handle prey, or on prey s i z e or prey type depending upon the p a r t i c u l a r t i n t i n n i d species. 'Negative' s e l e c t i o n of some types of laboratory phytoplankton i n mixed-prey samples was also shown f o r some t i n t i n n i d species, p a r t i c u l a r l y Tintinnopsis subacuta on members of the Cryptophyceae. Feeding rates measured with the accumulation method were equivalent i i to 0.65% m l / h r / t i n t i n n i d f o r T_. subacuta and usually much l e s s . Feeding rates f o r t h i s species measured with the Coulter Counter technique ranged from 0.33 to 3.8% m l / h r / t i n t i n n i d . Very l i t t l e feeding was observed d i r e c t l y but feeding rates estimated with t h i s method were somewhat higher than those estimated f o r the same species from accumulation experiments. Ti n t i n n i d s apparently both consumed, and caused the production of p a r t i c l e s during experiments. Correlations between feeding rate and 9 other experimental v a r i a b l e s were such that i t would be impossible to predict the feeding rate of a t i n t i n n i d species using only the s i z e d i s -t r i b u t i o n of avaifeble p a r t i c u l a t e biomass of l e s s than 20 um diameter. There were large differences between the apparent feeding rate asymptotes of T_. subacuta and _S_. ventricosa as measured with the Coulter Counter and the accumulation method. The l a t t e r method gave lower asymptotes than d i d the former. Ivlev e l e c t i v i t y indices for T_. subacuta were most consistently p o s i t i v e i n those middle Coulter s i z e classes which also showed the greatest growth i n controls. Increased temperature had l i t t l e e f f e c t on the rate of food accumu-l a t i o n by four t i n t i n n i d species, but there was some evidence of a f a s t e r rate of disappearance of ingested food at very high temperatures. The r e l a t i o n s h i p between the gain of new food and the loss of old food i n i n -d i v i d u a l T. subacuta and Stenosomella ventricosa was highly v a r i a b l e and may strongly r e f l e c t the p h y s i o l o g i c a l h i s t o r y of the c e l l . The rate of gain of new food may be l a r g e l y independent of the amount of old food i n a t i n t i n n i d , but the average rate of loss of o l d food i s f a s t e r i n c e l l s given new food than i n starved c e l l s . i i i I t was shown that natural concentrations of T_. sub acuta can apparently con t r o l the growth of natural populations of phytoplankton of l e s s than 20 um d i a . i n under 24 hours. From a comparison with some other types of microzooplankton i t was concluded that the larger species of t i n t i n n i d could probably have a p o t e n t i a l l y predominant e f f e c t upon the highly pro-ductive phytoplankton of l e s s than 10 um diameter i n English Bay and other coastal l o c a l i t i e s . i v TABLE OF CONTENTS Page ABSTRACT i TABLE OF CONTENTS i v LIST OF TABLES v i LIST OF FIGURES i x ACKNOWLEDGEMENTS x i 1. INTRODUCTION 1 2. TINTINNID BIOLOGY . 5 3. MATERIALS AND METHODS a) Sampling and i n i t i a l treatment 32 b) Experimental methods (i) General comments 35 ( i i ) Counts of accumulated food 38 ( i i i ) Observations of feeding behaviour 39 (iv) Coulter Counter experiments 41 Glossary 49 4. RESULTS AND DISCUSSION a) Accumulation experiments (i) Q u a l i t a t i v e r e s u l t s 51 ( i i ) Quantitative r e s u l t s 60 b) Observations of t i n t i n n i d motions and feeding behaviour 117 c) The e f f e c t of microzooplankton on natural and laboratory phytoplankton populations (Coulter Counter experiments) 127 5. GENERAL DISCUSSION 163 Table of Contents (Cont'd) Page 6. SUMMARY 179 REFERENCES 182 APPENDICES 187 v i TABLE 1, LIST OF TABLES Page L i s t of t i n t i n n i d species and t h e i r measurements. 52 TABLE 2. Food eaten by microzooplankton. 56 TABLE 3. Eutintinnus tubulosus feeding on Isochrysis galbana at two concentrations. 62 TABLE 4. Eutintinnus tubulosus feeding on Monochrysis-l u t h e r i at two concentrations. 62 TABLE 5. Eutintinnus tubulosus feeding on Monochrysis l u t h e r i at three concentrations. 64 TABLE 6. Tintinnopsis parvula feeding on Monochrysis l u t h e r i at three concentrations. 64 TABLE 7. Tintinnopsis subacuta feeding on D u n a l i e l l a t e r t i o l e c t a at three temperatures and three food l e v e l s . 66 TABLE 8. Tintinnopsis parvula and Tintinnopsis c y l i n d r i c a feeding on Monochrysis l u t h e r i i n dim l i g h t and i n darkness. 68 TABLE 9. Eutintinnus tubulosus and Helicostomella k i l i e n s i s feeding on Monochrysis l u t h e r i at three concentrations 69 TABLE 10. Various t i n t i n n i d species feeding on 'new' and 'old' cultures of D u n a l i e l l a t e r t i o l e c t a at four concentrations. 71 TABLE 11. Tintinnopsis parvula feeding on Isoselmis ssp. and Monochrysis l u t h e r i . 73 TABLE 12. Tintinnopsis subacuta feeding on E t i t r e p t i e l l a sp. and Isochrysis galbana. 73 L i s t of Tables (cont'd) Tintinnopsis subacuta feeding on E u t r e p t i e l l a  sp.,Isochrysis galbana and D u n a l i e l l a t e r t i o l e c t a . Tintinnopsis subacuta and Tintinnidium  mucicola feeding on E u t r e p t i e l l a sp. and Isoselmis sp. Various t i n t i n n i d species feeding on Monochrysis l u t h e r i and D u n a l i e l l a t e r t i o l e c t a . Tintinnopsis subacuta and Stenosomella  ventricosa feeding on E u t r e p t i e l l a sp., Monochrysis l u t h e r i and Isoselmis sp. s i n g l y and i n combination. Tintinnopsis subacuta (etc.) starved f o r various periods i n f i l t e r e d seawater feeding on D u n a l i e l l a t e r t i o l e c t a at unknown, but dense, concentrations. Tintinnopsis subacuta and other predators starved f o r various periods and feeding on E u t r e p t i e l l a sp. Loss rate of Stenosomella n i v a l i s at two l e v e l s of d i l u t i o n of medium with f i l t e r e d seawater. Change of food contents of -T-intinnidium  mucicola with time at four l e v e l s of d i l u t i o n o of medium with f i l t e r e d seawater. Tintinnopsis subacuta, T_. parvula, _T. rapa and Tintinnidium mucicola feeding on new food - Monochrysis l u t h e r i and Cryptomonas sp., and l o s s rate of o l d food of various types at four temperatures. v i i i L i s t of Tables (cont'd) TABLE 22. Feeding and loss rates of Tintinnopsis subacuta; l o s i n g Monochrysis l u t h e r i and Plagioselmis sp. and e i t h e r starved or gaining E u t r e p t i e l l a sp. and Isoselmis sp. Page 99 TABLE 23. Accumulation and l o s s rates of Tintinnopsis c y l i n d r i c a , Helicostomella k i l i e n s i s , Tintinnidium mucicola and Eutintinnus l a t u s , feeding oh Monochrysis l u t h e r i or Isoselmis sp., or starved; and l o s i n g M. l u t h e r i , Isoselmis sp. or D u n a l i e l l a t e r t i o l e c t a . 101 TABLE 23A. Summary of accumulation experiments with Tintinnopsis subacuta. 103A TABLE 24. The r e l a t i o n s h i p between t i n t i n n i d c e l l length and number of accumulated food items i n two species taken from d i f f e r e n t experiments. 115 TABLE 25. The e f f e c t of immobilization by sonication on the successful ingestion of a l g a l f l a g e l -l a t e s by the t i n t i n n i d , Eutintinnus l a t u s . 121 TABLE 26. Observed contact rates of various t i n t i n n i d species on na t u r a l and laboratory food items. 124 TABLE 27. Mult i p l e c o r r e l a t i o n c o e f f i c i e n t s from Coulter Counter experiments. 129 TABLE 28. Microzooplankton lower threshold feeding values and regression c o e f f i c i e n t s of Logmean E (variable 3) when food consumption rate (variable 1) i s zero. 136 TABLE 29. Results of Coulter Counter experiments with Synchaeta l i t t o r a l i s and Synchaeta sp. eating D u n a l i e l l a t e r t i o l e c t a . 159 TABLE 30. Approximate r e l a t i v e s i z e s and feeding rates of various types of marine microzooplankton. 177 FIGURE 1. FIGURE 2. FIGURE 3. FIGURE 4. FIGURE 5. FIGURE 6. FIGURE 7. FIGURE 8. FIGURE 9. FIGURE 10. i x LIST OF FIGURES Page Diagram of Fa v e l l a sp. (modified from 7 Campbell, 1927). Possible t h e o r e t i c a l r e l a t i o n s h i p s be- 15 tween t i n t i n n i d l o r i c a length and frequency. L o r i c a length-frequency data f o r Tintinnopsis 18 subacuta from three successive f i e l d samples. Seasonal abundance and l o r i c a lengths of 100 20 Paraf a v e l l a denticulata.= Relationship between t i n t i n n i d c e l l volume 53 and maximum observed volume of i n d i v i d u a l food items. Relationship between the volume of old food an 108 and new food contained by i n d i v i d u a l T i n -tinnopsis^ subacuta. Relationship between the volume of old food 110 and new food contained by i n d i v i d u a l T i n -tinnopsis subacuta and Stenosomella ventricosa. Relationship between e l e c t i v i t y values of 140 Tintinnopsis subacuta on natural p a r t i c l e s and the mean diameter of Coulter Counter s i z e classes.. Relationship between e l e c t i v i t y values of 143 Tintinnopsis subacuta on natural p a r t i c l e s and the Logmean E values of each Coulter Counter s i z e c l a s s . Relationship between the e l e c t i v i t y values 145 of Tintinnopsis subacuta on natural p a r t i c l e s and the t o t a l Logmean E values of a l l Coulter Counter si z e classes. X L i s t of Figures (cont'd) Page FIGURE 11. Relationship between the e l e c t i v i t y values 148 of Tintinnopsis subacuta on natural par-t i c l e s and the changes i n control values (C 0/C ) i n each Coulter Counter s i z e c l a s s . FIGURE 12. Relationship between the changes i n the 152 t o t a l p a r t i c l e volume of Coulter Counter control (C^/C^) and experimental (E^/E^) containers at various concentrations per ml. of Tintinnopsis subacuta and J_. parvula on natural p a r t i c l e s . FIGURE 13. Relationship between the changes i n the 154 t o t a l p a r t i c l e volume of Counter Counter control (C2/C^) and experimental (E^/E ) containers at various concentrations or Tintinnopsis subacuta per ml. on laboratory food. FIGURE 14. Relationship between the changes i n the 156 t o t a l p a r t i c l e volume of Coulter Counter control (C2/C^) and experimental (E^/E ) containers at various concentrations or Stenosomella ventricosa and Barnacle and copepod n a u p l i i on laboratory food and Barnacle and copepod n a u p l i i on natural p a r t i c l e s . x i ACKNOWLEDGEMENTS I thank Drs. P.A. Lark i n , J.D. Berger, T.R. Parsons and F.J.R. Taylor for advice and assistance during t h i s study. I am g r a t e f u l for f i n a n c i a l assistance during the project from Dr. B. McK. Bary and Dr. T.R. Parsons, and the work could not have been completed without generous f i n a n c i a l help from Dr. P.A. Larkin. Dr. E.S. G i l f i l l a n and Mr. T. Gossard were of great help with advice on s t a t i s t i c a l and computing problems. I am very glad to acknowledge the laboratory f a c i l i t i e s and equipment provided by Drs. Bary, Parsons, Taylor and Dr. A.G. Lewis and the s t a f f of the P a c i f i c B i o l o g i c a l Station, Nanaimo. I g r a t e f u l l y thank Mrs. R. Waters for providing some phytoplankton cultures and media. Drs. J.D. Berger, D.J. Rapport and F.J.R. Taylor k i n d l y allowed me to r e f e r to t h e i r unpublished work. The help of Miss H. Halm and Mrs. M.E. Newell was e s s e n t i a l to the preparation of t h i s t h e s i s . Most of a l l my g r a t e f u l thanks go to my wife Janice, for sustaining me throughout t h i s work. 1. I) INTRODUCTION This study consists of a l a r g e l y experimental i n v e s t i g a t i o n of the feeding a b i l i t i e s of the large and common c i l i a t e s , p a r t i c u l a r l y the t i n -t i n n i d s , from the marine plankton near Vancouver. Our knowledge of the structure and i n t e r a c t i o n s of marine planktonic food webs i s poor; t h i s ignorance i s unfortunate i n view of the large d i s -crepancies between various estimates of p o t e n t i a l t o t a l oceanic f i s h production, such as for example those taken from data on catches of f i s h , and from measurements of marine primary p r o d u c t i v i t y (Steele, 1965). One of the areas of greatest ignorance includes the food l i n k s to and from microzooplankton. C i l i a t e protozoa are the smallest and most abundant of the microzooplankton (30 - 1000 urn) organisms i n most oceans. In some marine areas, c i l i a t e s also form an important part of the t o t a l microzoo-plankton biomass, and (presumably) metabolic a c t i v i t y (Beers and Stewart, 1969B; Zaika and Averina, 1969; Zenkevitch, 1963). T i n t i n n i d s are among the l a r g e s t of the c i l i a t e microzooplankton, but almost nothing i s known of t h e i r ecology i n the sea or i n freshwater. Since much of the t o t a l m o r t ality i n a cohort i n many species of f i s h apparently occurs from st a r v a t i o n at a very early age, a great increase i n information on the tropho-dynamics of t h e i r p o t e n t i a l prey (often small microzooplankton) mayybe v i t a l to any understanding of the large natural yearly v a r i a t i o n s i n s u r v i v a l of cohorts of f i s h (Korniyenko, 1971). Furthermore, LeBrasseur and Kennedy (1972) think that i n v e s t i g a t i o n of the feeding habits and rates of production of very small (<50 urn) c i l i a t e s w i l l be required to a i d i n estimating the v i t a l winter n u t r i t i o n arid development 2 of the dominant macrozooplankters of the open subarctic P a c i f i c Ocean. The l a t t e r are c r u c i a l to the food-webs of the adults of several species of f i s h of commercial importance, e.g. sockeye s a l m o n S i m i l a r l y , Eggert (1973) states that t i n t i n n i d s are very numerous i n Lake Baikal and are im-portant to the winter n u t r i t i o n of copepod n a u p l i i i n that lake. The importance of microzooplankton to t h e i r predators at l e a s t p a r t l y depends upon the rates and e f f i c i e n c i e s with which they incorporate the nannoplankton which i s unavailable to the la r g e r organisms. Subjective f i e l d observations and estimates of microzooplankton feeding and producti-v i t y must be he a v i l y augmented with experimental work,but the feeding mechanisms of diverse microzooplankters cannot simply be extrapolated from those more e a s i l y investigated i n crustacean macrozooplankton. Although the crustacean members of the microzooplankton can be expected to feed l a r g e l y by f i l t e r - f e e d i n g with a f a i r l y r i g i d ' f i l t e r - b a s k e t ' , as do the larger and frequently studied crustacean macrozooplankton (Poulet, 1973), many of the microzooplankton including the t i n t i n n i d s and other c i l i a t e s , use c i l i a to obtain i n d i v i d u a l food items and th i s s p e c i a l i z e d mode of feeding should be investigated i n i t s own r i g h t . The papers of Strathmann (1971) and Strathmann, et. a l . (1972) represent most of what i s known of the feeding behaviour of some t y p i c a l riearshore metazoan planktonic c i l i a r y feeders, i n c l u d i n g echinoderm l a r v a e . Nothing i s known of the p r o d u c t i v i t y or feeding rates of t i n t i n n i d s i n s i t u . There i s also no published quantitative information on the feeding a b i l i t i e s of t i n t i n n i d s i n the laboratory and l i t t l e more i s known of t h e i r growth rates i n laboratory c u l t i v a t i o n (Gold, 1971, 1973). Indeed, very l i t t l e i s known of the feeding a b i l i t i e s of any marine c i l i a t e , benthic or 3 planktonic from which one might estimate the feeding rates to be expected from t i n t i n n i d s . Pavlovskaya (1973) has discussed the feeding and growth of a few c i l i a t e s found i n the l i t t o r a l zone of the Black Sea, and Fenchel (1968) has studied the food contents and reproductive rates of some benthic c i l i a t e s from the B a l t i c Sea. Hamilton and Preslan (1969) have grown one species of a small bacteria-grazing planktonic c i l i a t e i n cu l t u r e . There i s rather more, but s t i l l fragmentary data on some freshwater species. For example, Klekowski et. al.-(1972) gives some d e t a i l s of the energy budget of one species of benthic freshwater c i l i a t e and Goulder (1972, 1973) has i n d i r e c t l y estimated the feeding rate of a c i l i a t e free-swimming i n a pond. In general, the smaller the organism the smaller the absolute amount of food i t removes per unit time, and the fas t e r i t s reproductive rate. However, as c i l i a t e s u s ually greatly outnumber other microzooplankton and macrozooplankton; the t o t a l feeding e f f e c t of c i l i a t e s per unit volume of seawater may, under some circumstances, provide s i g n i f i c a n t competition i n removing phytoplankton, f o r macrozooplankton such as large copepods and euphausiids. I t has been shown that some copepods and euphausiids have a lower si z e threshold i n t h e i r feeding below which they graze phytoplankton poorly, i f at a l l (Parsons and LeBrasseur, 1970). I t i s of i n t e r e s t to determine whether t i n t i n n i d s and other c i l i a t e s would u t i l i z e these very small (2-15 um) food items. In order to obtain i n i t i a l answers to the problem of the l i k e l y e f f e c t s of t i n t i n n i d s as predators and competitors t h i s study was intended to be a broad and general experimental i n v e s t i g a t i o n of feeding rates and preferences i n the larger l o c a l species. More s p e c i f i c a l l y , the i n t e n t i o n was: 1. To investigate the range of material eaten by t i n t i n n i d s and i t s over-lap with the food of la r g e r zooplankton. 2. To obtain measurements of the feeding rates of t i n t i n n i d c i l i a t e s and other microzooplankton on natural types of food. 3. To make such measurements with more than one technique and i n such a way (e.g. by s i z e separation of natural zooplankton samples and by using a Coulter counter) that as f a r as possible comparisons could be made between d i f f e r e n t species of t i n t i n n i d s and between t i n t i n n i d s and other microzoo-plankton. 4. To inves t i g a t e the circumstances under which prey s e l e c t i o n ( i f any) occurs i n various t i n t i n n i d species. 5. To examine p a r t i c u l a r l y the e f f e c t of such factors as food concentrations prey s i z e , hunger state and'temperature upon t i n t i n n i d feeding rates. 6 . If pos s i b l e , to in v e s t i g a t e the r e l a t i o n s h i p between feeding r a t e , digest ion r a t e , and growth rate of t i n t i n n i d s on phytoplankton cultured from natural samples; and to compare t h i s information with observations made during natural t i n t i n n i d 'blooms'. 7. To estimate the p o t e n t i a l a b i l i t y of t i n t i n n i d s to control the growth of populations of small phytoplankton c e l l s . 5 2) TINTINNID BIOLOGY General The major emphasis of t h i s study has been on attempts to obtain q u a l i -t a t i v e and quantitative information on the feeding rates of several of the l o c a l t i n t i n n i d species. During t h i s work, much information (often q u a l i -t a t i v e ) on various aspects of t i n t i n n i d biology has accrued. Since the natural h i s t o r y of t i n t i n n i d s i s generally so poorly known, some d e t a i l s from the l i t e r a t u r e (with references) plus some of my observations are presented here. Various aspects of t i n t i n n i d biology have been treated i n the following papers: Systematics — Marshall (1969); Tappan and Loeblich (1968); Z e i t z s c h e l (1969) and many others. D i s t r i b u t i o n of species — Many papers quoted i n Z e i t z s c h e l (1969). D i s t r i b u t i o n of numbers and biomass — Beers and Stewart (1969B,11971); Z e i t z s c h e l (1969) ; Zenkevitch (1963). Seasonal and v e r t i c a l d i s t r i b u t i o n — Beers and Stewart! (1969A) ; V i t i e l l o (1964); Zaika (1972). V e r t i c a l migration — Eggert (1973); r Zaika and Ostrovskaya (1972). Seasonal v a r i a t i o n i n numbers and s i z e — Burkovsky (1973). Growth i n culture — Gold (1971, 1973). Morphology of L o r i c a — Biernacka (1965) ; Halme and Lukkarinen (1960); Burkovsky (1973). Cytology — Campbell (1926, 1927). U l t r a s t r u c t u r e — Laval (1971, 1972, 1973). T i n t i n n i d s form an Order i n the Subclass S p i r o t r i c h a which also includes the c l o s e l y r e l a t e d Order O l i g o t r i c h i d a and others of the more hig h l y evolved c i l i a t e s i n the Subphylum Cil i o p h o r a . Most no n - t i n t i n n i d marine planktonic 6 c i l i a t e species are O l i g o t r i c h s . Both orders are characterized by possess-ion of few somatic c i l i a and an an t e r i o r o r a l opening surrounded by a large complex c i l i a r y structure which serves both to move, and provide food f o r the organism. T i n t i n n i d s are attached p o s t e r i o r l y by a t h i n c o n t r a c t i l e structure, to the side or bottom of an external sheath (or l o r i c a ) of carbo-hydrate or proteinaceous material secreted by the c e l l . This l o r i c a , which may be e i t h e r transparent, or nearly opaque and covered with fo r e i g n p a r t i -c l e s , normally covers the posterior f o u r - f i f t h s of the c e l l and may be many times longer than the l a t t e r . The feeding (or adoral) c i l i a are normally completely exposed and project at an angle l a t e r a l l y beyond the l o r i c a when the organism i s extended. (see F i g . 1).. O l i g o t r i c h c i l i a t e s possess-no l o r i c a , but some have a t i g h t l y - f i t t i n g p osterior sheath of carbohydrate material and they generally have proportionately larger o r a l c i l i a than do t i n t i n n i d s . Food items are drawn towards the c i l i a t e by means of a vortex created by the anterior c i l i a , as the c e l l rotates and moves i n t e r m i t t e n t l y i n a h e l i c a l path of a shape c h a r a c t e r i s t i c f o r each species. Unlike some of the O l i g o t r i c h s , the l o c a l t i n t i n n i d s are incapable of making large and sudden 'jumping' movements. There i s some preliminary sorting of food i n the anterior c i l i a r y region and mucus may perhaos be produced to aid i n the capture of food (Laval, 1972). I n t r a c e l l u l a r d i g e s t i o n follows the phago-cyt o s i s of i n d i v i d u a l food items. Egestion of the undigested material as ei t h e r s i n g l e objects or as clumps occurs at some posterior s i t e i n the c e l l plasma membrane. Most t i n t i n n i d s must then remove the egested material from i n s i d e the l o r i c a . This i s done by a row of t h i n l a t e r a l c i l i a which pass the egesta along the narrow space between c e l l and l o r i c a wall and out over the 7 Fig. 1. Diagram of FAVELLA sp. (modified from Campbel l , 1927) showing adoral c i l ia , a.c. • anus, an., cytopharynx , cyt, food vacuoles, f.v., lorica, lor. , macro-nucleus, macron., micronucleus, micron., oral plug, o.p., peduncle, ped. 8 l i p of the l o r i c a . In the genus Eutintinnus, egested material remains post-e r i o r to the c e l l and may eventually pass out of the l o r i c a through the p o s t e r i o r opening. Ti n t i n n i d s are generally most abundant i n the upper part of the euphotic zone and are nearly always more abundant i n inshore waters than offshore. (Beers and Stewart 1969B, 1971; Z e i t z s c h e l 1969). There i s evidence that some species make short d i u r n a l verfc£<sali migrations away from the surface at dawn and towards the surface at dusk, (Eggert, 1973; Zaika and Ostrovskaya, 1972) . Taxonomy The taxonomic designations of Marshall (1969) have been followed as c l o s e l y as possible i n t h i s study, although even t h i s p r a c t i c e has involved the a r b i t r a r y choice of s p e c i f i c name i n some cases. This problem w i l l be dealt with again i n the comments on l o r i c a morphology l a t e r i n t h i s Section. Seven genera of t i n t i n n i d s represented by t h i r t e e n species have been studied i n the course of t h i s p r o j ect. A few other species occur i n Georgia Straight but f o r these no data has been obtained. Ten of these species are commonly found throughout the year, often i n the same sample. There i s an e i g h t - f o l d range of c e l l length and a three hundred-fold range of c e l l volume between the smallest species Tintinnopsis nana and the l a r g e s t , F a v e l l a  serrata (see Table 1). A s i n g l e inshore microzooplankton sample has been found to contain up to s i x species of other phagotrophic c i l i a t e s , up to three species of r o t i -f e r s , and larvae and adults of three or four species of small copepods. More temporary microzooplankters include the l a r v a l forms of barnacles, 9 bryozoans, polychaetes, and bivalve and gastropod molluscs. Temperature and s a l i n i t y e f f e c t s The bulk of the samples were taken by sampling from shore i n English Bay and Coal Harbour, Vancouver. Samples were also taken from Boundary Bay, Horseshoe Bay, c e n t r a l Georgia S t r a i t , and from Departure Bay, Saanich I n l e t , and from harbours at Sydney, V i c t o r i a and Tofino on Vancouver Island. No species was found to be unique to any p a r t i c u l a r area, and t h i s would almost c e r t a i n l y s t i l l be true on a much broader geographic scale i n inshore waters. Also, as expected i n a semi-estuarine area, a l l species were extremely t o l -erant to slow changes i n temperature, and to f a s t or slow changes over a considerable range i n s a l i n i t y . These a b i l i t i e s were checked i n b r i e f exper-iments. Most of the t i n t i n n i d species were found at a l l the various natural combinations of temperature and s a l i n i t y , but large numbers or 'blooms' of f o r example, 2 or more individuals/ml generally occurred i n water of some le s s extreme combination of temperature and s a l i n i t y . The seasonal trends of temperature and s a l i n i t y of the l o c a l surface water are inversely cor-r e l a t e d , due to the great influence i n mid-summer of the Fraser River freshet. 'Blooms' of various species occurred at any time from March to December, usually a f t e r a few days of f i n e weather, but were most frequent i n April-May and September-November. Those t i n t i n n i d species which occurred i n large numbers ('blooms') (1-15/ml) p r i m a r i l y i n the months March-May and September-December were Tintinnopsis subacuta, T_. parvula, T_. rapa, Stenosomella ventricosa and S^. nivalis;. In those months surface temperatures i n English Bay are between 8 and 15° C and s a l i n i t i e s are between 10 and 27%0 . Helicostomella  k i l i e n s i s and Eutintinnus tubulosus were nnumeWo"*!© only between May and 10 September, when surface water temperatures are between 12 and 20°C. Tintinnopsis nana and Tintinnidium mucicola have been found i n large numbers ffinnevery month, but January and February. Tintinnopsis c y l i n d r i c a was at i t s most abundant from May to September, but was never as numerous as 1/ml. I t i s curious that T_. c y l i n d r i c a (frequently) and one i n d i v i d u a l of T_. subacuta, were the only t i n t i n n i d species seen to contain various stages of an unidenti-f i e d d i n o f l a g e l l a t e i n t e r n a l p a r a s i t e . The other three species i n Table 1 (section 4) were never numerous. Eutiritiririus l atus was seen only i n summer, and Fa v e l l a serrata and Ptychocyclis acuta (very rare) were seen i n l a t e summer-fall i n waters of a s a l i n i t y exceeding 15%>. However, i t i s l i k e l y that a l l other environmental factors are of secondary importance to a good supply of su i t a b l e food i n the production of large numbers of t i n t i n n i d s . L o r i c a morphology One t i n t i n n i d species, Tintinnidium mucicola, appears to have a l o r i c a to which p a r t i c u l a t e matter may adhere at any time. Others such as those i n the genera Tintinnopsis and Stenosomella have l o r i c a s which appear to be 'sticky ' f o r a short time only a f t e r formation and are l e s s so than that of Tintinnidium mucicola even then. Species of the genera F a v e l l a , Ptychocyclis, Helicostomella and Eutintinnus i n l o c a l waters never have p a r t i c l e s adhering to t h e i r l o r i c a s even when inwater with much small d e t r i t u s , although the -l o r i c a of E. tubulosus has been seen to be covered by a coating of f i n e d e t r i t u s which could be e a s i l y discarded. Several species of the genera T intin n o p s i s and Stenosomella can be kept i n laboratory conditions f o r short periods where they usually form apparently normal l o r i c a s with no adhering d e t r i t u s whatever. The function ( i f any) of d e t r i t u s on l o r i c a s i s not obvious. 11 L o r i c a 'repair' i s thought to be c a r r i e d out by some t i n t i n n i d species (Biernacka, 1965) and occasional i r r e g u l a r i t i e s i n l o r i c a morphology may be due to imperfect r e p a i r following 'accidents'. The shape of the l o r i c a s of t i n t i n n i d s i n laboratory cultures i s o c c a s i o n a l l y abnormal (Gold, 1971; and personal observation). This may be due to the lack of s u i t a b l e d e t r i t u s i n such c u l t u r e s , but abnormalities also occur i n cultures of Helicostomella  k i l i e n s i s ^ ( s e e above). Of more importance to the study of the taxonomy, palaeontology and pop-u l a t i o n dynamics of t i n t i n n i d s i s the v a r i a t i o n i n the length of l o r i c a s w ithin a species. T i n t i n n i d s are the only c i l i a t e s which appear to have l e f t f o s s i l s (Tappan and Loeblich, 1968) due to the nature of the p a r t i c l e s adhering to the l o r i c a s of some species. At present, the taxonomy of t i n -t i n n i d s i s based e n t i r e l y on the morphology of the l o r i c a , and species are designated l a r g e l y upon the basis of l o r i c a length (e.g. Marshall, 1969). Marshall, i n p a r t i c u l a r , recognized t h i s state of a f f a i r s to be unsatisfactory. For example, Burkovsky (1973) has suggested from samples c o l l e c t e d over two years that eleven species i n the genus Par a f a v e l l a from the White Sea are a l l v ariants of P_. d e n t i c u l a t a . I t has been observed i n t h i s study that l o r i c a length i s at l e a s t partly-a function of recent environmental conditions; and that the mean length, and the greatest length, of the l o r i c a s i n one apparent population of a species may d i f f e r considerably i n two samples taken a few days apart. In general i n t h i s area l o r i c a s tend to be longer i n conditions of p l e n t i f u l food, and t h i s i s often noticeable near the end of 'blooms'. However, c e l l s with p a r t i c u l a r l y long l o r i c a s are someimes found i n samples i n which there are few i n d i v i d u a l s of that species. Burkovsky (1973) found a generally 12 inverse r e l a t i o n s h i p between the l o r i c a lengths of Parafavella denticulata and phytoplankton concentrations and temperature. The diameters of l o r i c a s are much l e s s v a r i a b l e than l o r i c a lengths (this study and Burkovsky, 1973). The only completely r e l i a b l e diagnostic features .in' a genus of t i n t i n n i d s such as Tintinnopsis appear to be those most v i s i b l e i n l i v e or very c a r e f u l l y preserved organisms. Such features might include the average c e l l length and diameter, and the length of the adoral c i l i a . Differences i n swimming and feeding behaviour are also a useful but subtle taxonomic guide (see Section 4B). Generally the c e l l s of most t i n t i n n i d species are approximately c y l i n -d r i c a l f o r the anterior-two-thirds of t h e i r lengths, and c o n i c a l for the posterior one-third. The most obvious exceptions to t h i s r u l e are Stenosomella  ventricosa and S^. n i v a l i s , where the c e l l has a short anterior c y l i n d r i c a l s ection, and i s sub^spherical p o s t e r i o r l y , as are the l o r i c a s of these species. During starvation the shape of the c e l l s of a l l species change greatly to something l i k e that of a cone, and the c e l l volume may decrease by more than 50%. C e l l s of t h i s shape are p a r t i c u l a r l y common i n winter. The r a t i o of c e l l length to l o r i c a length i s v a r i a b l e within a species fo r several reasons. Simple transverse binary f i s s i o n without sexual recom-bi n a t i o n i s by f a r the commonest form of t i n t i n n i d reproduction. At the end of t h i s process, the anterior daughter c e l l breaks o f f from the posterior daughter c e l l and moves away without a l o r i c a . Therefore, the anterior daughter possesses the 'parental' o r a l c i l i a , the posterior daughter possesses the parental l o r i c a , and they share the parental cytoplasm and assimilated food. The duration of the process of d i v i s i o n i s unknown but l a s t s at l e a s t for several hours ( i n most species). The daughter c e l l s are i n i t i a l l y h a l f 13 that of the maximum parental c e l l length, which generally seems to be a char-a c t e r i s t i c of each species, but which may vary somewhat with r e l a t i v e l y long-term environmental conditions and with the p h y s i o l o g i c a l capacities of a clone of c e l l s . Each daughter c e l l contains a random f r a c t i o n of the recently ingested food. I t i s not c e r t a i n to what extent the posterior daughter c e l l can continue to add to the length of the parental l o r i c a . Since l o r i c a s with recent additions at the o r a l end are f a i r l y common, e s p e c i a l l y on c e l l s i n laboratory cultures, l o r i c a addition may be possible f o r any 'young' c e l l . C e r t a i n l y , as the anterior daughter c e l l grows toward the maximum c e l l length, i t concurrently manufactures a complete new l o r i c a from the secretion of material held i n granules i n the anterior portion of the c e l l . The new l o r i c a i n v a r i a b l y hardens and grows from the posterior (or aboral) end of the new c e l l , often from a template c o n s i s t i n g of a small p a r t i c l e of d e t r i t u s . I t i s not known how c l o s e l y the rates of growth of c e l l and l o r i c a are r e l a t e d . The anterior daughter c e l l begins to feed very soon a f t e r d i v i s i o n and forms a complete.lorica r a p i d l y , usually i n a day or so. Whether the ultimate s i z e of the new l o r i c a i s c h i e f l y dependent on the n u t r i t i o n a l state of (a) the growing c e l l , or (b) the parental c e l l i s not c e r t a i n . When c e l l s of near maximum s i z e are seen in s i d e very small l o r i c a s , the cause i s probably some sudden stress r e s u l t i n g i n the detachment of the c e l l from i t s o r i g i n a l l o r i c a . A l t e r n a t i v e l y , t h i s condition may r e s u l t from some imbalance i n the r e l a t i v e rates of growth of c e l l and accretion of l o r i c a , perhaps over several generations. Short c e l l s with r e l a t i v e l y large l o r i c a s are eit h e r the r e s u l t of recent c e l l d i v i s i o n or, and l e s s l i k e l y , of starvation. A newly divided c e l l starved since d i v i s i o n would be both short and 't h i n ' . Some c i l i a t e s grown i n laboratory culture can be 10 to 20 - f o l d l a rger at high growth rates than at low growth rates due to 14 a delay i n the maximum rate of reproduction (Canale, e_t,al_. , 1973; Hamilton and Preslan, 1969). Therefore, comparisons between t i n t i n n i d c e l l s i z e s with-i n a species should best await further data on growth rates from laboratory cultures. Data c o n s i s t i n g of length-frequency d i s t r i b u t i o n s of t i n t i n n i d l o r i c a s taken from f i e l d samples, may give information about some aspects of the 'population dynamics' of the l o r i c a s ; and perhaps on the recent h i s t o r y of growth of the (semi-immortal) c e l l s themselves.. This sort of information can be obtained from almost no other f r e e - l i v i n g protozoa. Haime and Lukkarinen (1960) and Burkovsky (1973) presented the two previous analyses of t i n t i n n i d l o r i c a measurements from f i e l d samples. L o r i c a length-frequency data from a time-series of samples taken at very short i n t e r v a l s , may help to d i s t i n g u i s h between two extreme theories of l o r i c a a c c r e t i o n , assuming synchronous reproduction and given c e r t a i n as yet u n v e r i f i a b l e assumptions about l o r i c a 'mortality'. These theories are as follows: a) Lorica accretion i s coohtinuous throughout t h e ' l i f e ' of the contained c e l l and i s dependent upon the current n u t r i t i o n a l state of that c e l l ; and b) Lo r i c a accretion i s discontinuous and confined to new l o r i c a s , occurs only immediately following d i v i s i o n , and i s dependent upon the n u t r i t i o n a l state of the parental c e l l . If theory a) holds, then length-frequency diagrams of the l o r i c a s of one pop-u l a t i o n of one species from a time-series of consecutive samples 1 to 4, with conditions f o r growth improving i n that order and no l o r i c a m o r t a lity, would appear as i n Figure 2a); and i f theory b) holds, then the length-frequency diagrams would appear as i n Figure 2b)<. Intermediate patterns are possib l e . Biernacka (1965) b e l i e v e s , and I concur, that while most l o r i c a a c c r e t i o n occurs immediately a f t e r c e l l d i v i s i o n , i t can occur i n some species 15 Figure 2. Possible t h e o r e t i c a l r e l a t i o n s h i p s between t i n t i n n i d l o r i c a length and frequency (see text for d e t a i l s ) . Fig. 2 a) LORICA LENGTH GENETIC MAX. 2 b) GENETIC LORICA GENETIC MIN. LENGTH MAX. 1.7 at any time. Length-frequency diagrams of the l o r i c a s of one species of t i n t i n n i d from English Bay are shown i n Figure 3. Data from Burkovsky (1973) are shown i n Figure 4. These diagrams seem to in d i c a t e that theory b) i s a better des-c r i p t i o n of l o r i c a accretion than theory a) at l e a s t under some circumstances. There have been no previous t h e o r e t i c a l attempts at analyses of t h i s kind. It would be useful i n some future study to check these tentative ideas and other theories of l o r i c a growth and s i z e and age-dependent ( l o r i c a ) mor-t a l i t y r a t e s , etc., with data from a much wider range of samples and environ-mental conditions. I t i s probable that a new l o r i c a can be considered mainly as a metabolic 'cost' to the parental c e l l , and as such may be r e l a t i v e l y greatest i n those species which have the l a r g e s t r a t i o of l o r i c a length to c e l l length. One l o c a l species i n p a r t i c u l a r , Helicostomella k i l i e n s i s , possesses a l o r i c a with markings which may be of some future use i n estimating rates of l o r i c a a c c r e t i o n , etc. These markings consist of c l o s e l y adjacent narrow s t r i p s of material l a i d down i n the shape of a h e l i x on top of the anterior portion of the l o r i c a . These 'annuli' are i n v a r i a b l y about 4 um apart but t h e i r number per l o r i c a i s greatly v a r i a b l e ; and they are not always most numerous upon the longest l o r i c a s of H. k i l i e n s i s . In short, the r e l a -tionships between the rates of t i n t i n n i d c e l l growth and d i v i s i o n and the rate of annulus formation ( i f any); and between the l a t t e r and the rate of accre-t i o n of the whole l o r i c a , are unknown. Asexual reproduction T i n t i n n i d c e l l s do not n e c e s s a r i l y grow and divide most r a p i d l y when they contain the l a r g e s t amounts of recently ingested, or of assimilated food 18 Figure 3. Lorica length-frequency data for Tintinnopsis subacuta from three successive f i e l d samples. 25r 24 23 22 21 20 19 18 17 16 15 14 13 12 11 10 9 NOS.8 7 6 5 4. 3 2 1 19 n. n , n Unlh. 0.8 ML. of NET HAUL 30/4/70 20 30 40 50 60 70 80 90 100 110 120 5 4 NOS.3 2 1 20 •iff rl 30 40 50 60 70 25 ML. of UNCONCENTRATED SAMPLE 30/4/70 0.6/ML. n. n 80 90 100 110 120 8 . 7 6 5 NOS. 4 3 2 1 20 25 ML. of UNCONCENTRATED SAMPLE 3/5/70 n 1.6/ML. n HI n 30 40 50 60 70 80 90 100 110 120 7 6 NOS.J: 3 2 1 n . n n n . 25 ML. of UNCONCENTRATED SAMPLE 6/5/70 0.6/ML. n n n 20 30 40 50 60 70 80 90 LORICA LENGTH (um) 100 110 120 20 Figure 4. Seasonal abundance and l o r i c a lengths of 100 Par a f a v e l l a d e n t i c u l a t a . Data from Burkovsky (1973). MONTH NOS./j STN 1 MONTH NOS./ M -JULY 60 SEPT. 3000 JAN. MAY 200 170 180 190 200 210 220 230 240 250 260 270 280 290 300 310 320 330 340 350 360 370 380 3W 400 LORICA LENGTHS (ym) 22 (see Section 4)• Temperature may have a greater e f f e c t on d i v i s i o n rate than on feeding rate. The o r a l structures of the posterior daughter c e l l develop s p i r a l l y (from a primordium or anlage) over a period of several hours at a mid-point i n the l a t e r a l surface of the parental c e l l , which i s at that time probably close to i t s maximum length for that species (Campbell, 1926). The diameter of the new (posterior) o r a l region, and the length of the o r a l c i l i a around i t , increase together, as the anterior daughter c e l l i s extended forward well c l e a r of the l o r i c a . The l a t t e r i s f i n a l l y connected to the posterior daughter only by a t h i n strand of cytoplasm close to the new f u l l -sized o r a l region. Throughout at l e a s t the f i r s t part of c e l l d i v i s i o n the parental o r a l region remains s u p e r f i c i a l l y unchanged, (unlike that i n many groups of c i l i a t e s ) and may gather food. The eventual decline of v i a b i l i t y of an asexual clone (as shown by small s i z e , slow reproductive rate, etc.) i s a well known genetic phenomenon i n c i l i a t e s i n laboratory culture. This decline can only be arrested by sexual recombination. Presumably natural clones of t i n t i n n i d s are prone to such de-c l i n e s , but there i s no data on the subject. Sexual Reorganization This i s a much l e s s common phenomenon than asexual reproduction i n t i n -t i n n i d s , as i s the case i n a l l natural populations of c i l i a t e s . Mating i n -volves the p a i r i n g (or conjugation) of c e l l s belonging to some of the d i f f e r -ent mating types within a morphologically defined species of c i l i a t e f o r the purpose of genetic recombination through the exchange of micro n u c l e i . Conjiigants eventually form cytoplasmic connections i n the o r a l region for t h i s exchange. Conjugation has been seen most commonly i n t h i s study i n f a i r -l y abundant l a t e summer-fall populations of t i n t i n n i d s . For example, i n one 23 sample i n September 1973, approximately 25% of i n d i v i d u a l s of Stenosomella  n i v a l i s , about 20% of Tintinnopsis parvula and about 10% of T_. subacuta were i n conjugating p a i r s . The s p e c i f i c f a c t o r s which stimulate t h i s process are unknown. During a feeding experiment at that time, one p a i r of c e l l s of T\ subacuta remained i n conjugation for at l e a s t 1% hours and probably much longer, and d i d not feed. Nothing i s known of the d e t a i l s of the mating pro-cesses nor of the possible number of 'mating types' i n any species of t i n -t i n n i d . Gold and Pollingher (1971) claim to have observed an unusual type of sexual process i n a laboratory culture of one species of t i n t i n n i d involving a mobile microgamete which attached i t s e l f to the posterior end of the macro-gamete. Their evidence seems i n s u f f i c i e n t for t h e i r claim to be supported unreservedly. The phenomenon has not been seen i n t h i s study. Motion and Metabolism Throndsen (1973) has shown that i n a wide v a r i e t y of marine a l g a l f l a g -e l l a t e s there i s no obvious r e l a t i o n between c e l l s i z e and swimming rate and the distances t r a v e l l e d per second may range from 10 to 40 c e l l lengths i n d i f f e r e n t species. Rotation and gyration are the r u l e i n f l a g e l l a t e s but each species has a c h a r a c t e r i s t i c motion. Bullington (1925) studied the motion of c i l i a t e s and found that c e l l r o t a t i o n and h e l i c a l paths were char-a c t e r i s t i c of many species. He found that larger species were generally f a s t e r than small ones, and that the f a s t e r t h e i r speed the fewer the turns they made. Bullington also showed that most c i l i a t e species had a speed of 5 or 6 c e l l lengths per second, rather slower i n r e l a t i o n to s i z e than the f l a g e l l a t e s mentioned above, but much f a s t e r inyum/second. In Bullington's r e s u l t s the paths of the f a s t e r c i l i a t e s described fewer s p i r a l s per body 24 length t r a v e l l e d , than did the slower species. He states Prorbdon marinus (200 x 100 urn) as moving at about 1000 um/second. In t h i s study, Prorodon sp. has been seen to move at between 5 and 12 c e l l lengths/second, with r o t a t i o n but very infrequent h e l i c a l motions. Bullington ( l oc. c i t . ) d i d not study t i n t i n n i d s . I t has been occasionally possible to make d i r e c t measurements of t i n t i n n i d speeds i n t h i s study. For example, the speed of Stenosomella  ventricosa has been estimated at d i f f e r e n t times at four and at eight body lengths per second. The amount of energy expended on motion by very small organisms seems to be a very small proportion of t h e i r t o t a l metabolic energy losses. Halfen and Castenholz (1971) state that the energy required to move a g l i d i n g blue-green bacterium cannot be more than 5% of the energy produced by oxidative phosphorylation. Likewise, Pavlova and Lanskaya (1969) found that i n f i v e species of Black Sea d i n o f l a g e l l a t e s the percentage of a c t i v e (motion) r e s -p i r a t i o n i n the t o t a l metabolism was about 1%. In the case of the much larger copepods, Vlymen (1970) estimated that extensive d a i l y v e r t i c a l mig-rat i o n s would require the equivalent of l e s s thaii 1% of the basic metabolic rate of these organisms. In a l l swimming organisms smaller than about 1 cm. i n length energy i s expended mostly to overcome the v i s c o s i t y of the medium rather than the i n e r t i a of the organism. Also, turbulent flow i s n o t created by small, slow organisms, and therefore energy need not be expended to over-come i t . It i s not l i k e l y that the constant motion of t i n t i n n i d s w i l l be an important part of t h e i r t o t a l metabolic a c t i v i t y , p a r t i c u l a r l y i n warm water, the v i s c o s i t y of which i s lower than that of cold water. A very rough estimate of the t o t a l oxygen consumption of Tintinnopsis  subacuta was made with the use of a d i f f e r e n t i a l microrespirometer as 25 described by Swift (1974). Five hundred t i n t i n n i d s of one species from a net.sample were placed i n the respirometer i n 3 ml of f i l t e r e d seawater at 15 C. This was thought to be a necessary but unnaturally high degree of crowding, but Gold (1973) has grown healthy populations of t i n t i n n i d s i n higher concentrations than t h i s . Since no (or very l i t t l e ) food was a v a i l a b l e , the r e s p i r a t i o n measured might be considered that of mobile but very crowded, slowly starving organisms. However, since only 38% of the t i n t i n n i d s were a l i v e a f t e r the 18% hr experimental period, the r e s p i r a t i o n measured may be p a r t l y the r e s u l t of b a c t e r i a l metabolism. If the l a t t e r p o s s i b i l i t y i s ignored, the t o t a l oxygen consumption was the equivalent of between 1.6 and -4 5.9 x 10 ul/hr/Tintinnopsis subacuta. This i s equivalent to a mean d a i l y consumption rate of 0.009 u l 0^/ T_. subacuta. There are no comparable data for t i n t i n n i d s or other planktonic c i l i a t e s and l i t t l e for other microzooplankton; but Doohan (1973) states that a female Brachionus p l i c a t i l i s r o t i f e r carrying three eggs u t i l i s e s about 0.14 u l 02/rotifer/day.. Petipa, and Marshall and Orr have measured the r e -s p i r a t i o n of marine copepod n a u p l i i . In t h e i r r e s u l t s (summarized with other work by Marshall, 1973) A c a r t i a c l a u s i i stage V and VI n a u p l i i used about 0.10 u l 02/nauplius/day; and Calanus finmarchicus stage III n a u p l i i used about 2 0.19 u l 0 /nauplius/day. The metabolic rates for the adults of these two copepod species were about 15 times greater than that of the quoted nauplius for A. c l a u s i i females and 50 times greater i n the case of CJ. finmarchicus ( i n Marshall, 1973). The weight of A. c l a u s i i stage V and VI n a u p l i i can be calculated from Marshall (1973) to be about 0.10 ug dry weight. From the data given by Theilacker and McMaster (1971) a large i n d i v i d u a l of Brachionus p l i c a t i l i s 26 6 3 would have a volume of 1 to 2 x 10 um . If a s p e c i f i c density of 1.0, and a wet weight to dry weight r a t i o of 10/1 are assumed, then a dried ]3. p l i c a t i l i s female would weigh about 0.15 ug. Therefore the r e s p i r a t i o n r a t e s , and dry weights are s i m i l a r f or A. c l a u s i i l a t e naupliar stages and 13. p l i c a t i l i s females. Using the above conversions Tintinnopsis subacuta 4 3 (7 x 10 um ) would have addry weight of about 0.007 ug. Thus these three microzooplankton organisms have an approximately.similar r e s p i r a t i o n rate per ug dry weight. The r e s p i r a t i o n rate of T_. subacuta may be calculated i n terms of ug 12 Carbon/tintinnid/day thus: —hO.009 x 2 2 ^ x 1.0 (respiratory quotient) = 0.0048 ugC/tintinnid/day. If a carbon/dry weight r a t i o of 0.5 i s assumed, then the mean r e s p i r a t i o n rate given i s equivalent to about 137% TC. subacuta body weight/day. This f i g u r e seems a l i t t l e high, but not unreasonable i n view of the f a c t that a t i n t i n n i d may e a s i l y consume the equivalent of 2-300% of i t s body weight i n food i n a day.(seeeSeneral Discussion). A r e s p i r a t o r y quotient of 0.8 may be more sui t a b l e than one of 1.0 i f l i p i d s are the chief r e s p i r a t o r y substrate (see Materials and Methods Section). If so then the r e s p i r a t i o n rate would be equivalent to about 110% T_. subacuta body weight/ day. Feeding This subject w i l l be discussed i n more d e t a i l . i n other sections. The predominant l i v i n g items i n the environmnet of t i n t i n n i d s , which are small enough to be eaten by them, are small (3 to 30 jam) naked u n i c e l l u l a r f l a g e l -l a t e d algae. As t i n t i n n i d s are, i n a general sense, unselective feeders, i t i s not s u r p r i s i n g that t h e i r food content i n general, r e f l e c t s t h i s prepon-derance of f l a g e l l a t e s . Occasionally, small diatoms, thecate d i n o f l a g e l l a t e s , 27 s i l i c o f l a g e l l a t e s , other t i n t i n n i d s , d e t r i t u s p a r t i c l e s or b a c t e r i a (usually on the l a t t e r ) form part of the c e l l contents. Blooms of one or more species of f l a g e l l a t e s are usually soon followed within 1 or 2 days, or are accom-panied by, blooms of one or two species of t i n t i n n i d s . Blooms of food c e l l s are not always the r e s u l t of photosynthesis. For example, i n l a t e December 1972, unusually high numbers (for the time of year) of Tintinnidium mucicola were seen i n samples from English Bay i n which there were very high numbers of a f l a g e l l a t e (or coccoid bacterium) 2 - 3 /im i n diameter, and not much el s e . During the previous ten days the cloud cover had been complete, and heavy r a i n had f a l l e n almost continuously. As a r e s u l t , some small l a n d s l i d e s had added much t u r b i d run-off to the sea i n the sampling area. The s h o r t - l i v e d f l a g e l l a t e (or b a c t e r i a l ) bloom was therefore almost c e r t a i n l y dependent on the uptake of allochthonous organic compounds i n the surface l o w - s a l i n i t y water. Occasionally, blooms of f l a g e l l a t e s are not f o l -lowed by large numbers of t i n t i n n i d s . Except f o r those cases i n which the f l a g e l l a t e was obviously too large f o r the a v a i l a b l e t i n t i n n i d species to ingest, as for a bloom of the d i n o f l a g e l l a t e Prorocentrum micans i n 1971, there seem no simples explanations for these 'unaccompanied' blooms of f l a g e l -l a t e s . The opposite s i t u a t i o n also occurred, where a sample contained r e l a t i v e l y many t i n t i n n i d s . In such cases, the t i n t i n n i d s contained few food items or storage granules, as ah i n d i c a t i o n of a recent low l e v e l of food intake. A sample taken the following day often contained many fewer t i n t i n n i d s . There-fore the previous sample either represented a) a food-poor environment i n which t i n t i n n i d s had been temporarily concentrated by water movements; or b) an environment i n which t i n t i n n i d reproduction had 'overshot' that of i t s 28 food, and where the food had as a r e s u l t become greatly depleted causing the t i n t i n n i d population to 'crash' from starvation. Such overshoots are possible because declines i n the reproductive rate of laboratory populations of c i l i -ates and other protozoa are known to lag behind declines i n feeding and growth rates; (Mitchison, 1971; Williams, 1971, 1972). Among the microzooplankton as a whole, some of the larval, forms appear to take l i t t l e or no food; but otherwise (even amongst the t i n t i n n i d s ) there i s much food 'overlap', and possibly also competition f o r food, i f the l a t t e r i s ever l i m i t i n g . Although the c i l i a t e s are almost always the most numerous microzooplankters, and have the f a s t e s t reproductive rates; some of the meta-zoans, such as the r o t i f e r Synchaeta l i t t o r a l i s contact and ingest a l g a l c e l l s at a f a s t e r rate than many c i l i a t e s , (see Section 4c), and also reproduce very r a p i d l y (personal observations) . Large populations of S_. l i t t o t a l i s occur i n inshore water of f a i r l y low s a l i n i t y (<10%^ and high temperature (>15°C); and as such almost c e r t a i n l y have a greater e f f e c t on the f l a g e l l a t e popu-l a t i o n s than do c i l i a t e s or any other microzooplankton. There i s no published information on the feeding of t h i s genus of r o t i f e r , and very l i t t l e further information has been added during t h i s study (but see Section 4c). However, information on laboratory feeding, growth and energy budgets of the brackish-water r o t i f e r Brachionus p l i c a t i l i s may be found i n the papers of Doohan (1973) and Theilacker and McMaster (1971). Predation on and Among T i n t i n n i d s Questions concerning the i d e n t i t i e s and a c t i v i t i e s of the predators of planktonic c i l i a t e s are important to an understanding of planktonic food webs but there are only the sc a n t i e s t answers. Of a l l these c i l i a t e s , only (the l o r i c a s e of) t i n t i n n i d s leave recognisable remains a f t e r mastication. This 29 ensures that c i l i a t e s w i l l only be found i n natural samples, except by chance, in s i d e those predators with some form of primary i n t r a c e l l u l a r d igestion (e.g. other protozoa), those with r e l a t i v e l y weak powers of mastication (e.g. r o t -i f e r s and chaetognaths), or those with no mastication and slow rates of extra-c e l l u l a r d i g e s t i o n (e.g. some f l a t f i s h l a r v a e ) . Other l i k e l y predators of t i n t i n n i d s would need to be checked with the use of r a d i o a c t i v e l y - l a b e l l e d tracer experiments. Data on the p o t e n t i a l i n g e s t i o n of t i n t i n n i d s has come from observation of the food grooves of benthic c r i n o i d s , and the mouthparts of planktonic crustacea (Bainbridge^1958). I t i s improbable that planktonic c i l i a t e s have any s p e c i a l i z e d predators, and l i k e l y predators would include any f a i r l y indiscriminate planktonic or benthic organisms well adapted f o r the ingestion of objects between 30yum and 300/am long. C i l i a t e s form about 90% of the d i e t of the larvae of three species of freshwater f i s h during the f i r s t four days of t h e i r exogenous mode of feed-ing (Korniyenko, 1971). As the motions of most c i l i a t e s are not e r r a t i c , they almost c e r t a i n l y do not e l i c i t searching movements by some r a p t o r i a l predators such as chaetognaths and copepods, but are probably captured and eaten as a r e s u l t of random encounters. However, Pearre (1973), has shown that t i n t i n n i d s can form up to 18% of the die t of young (Stage I) chaetognaths. In t h i s study, various t i n t i n n i d species have been seen inside polychaete larvae, the r o t i f e r Synachaeta l i t t o r a l i s , the large c i l i a t e s ProrOdon sp. and Strombidium (Lohmaniella) s p i r a l i s , and the t i n t i n n i d s Tintinnopsis  subacuta, T. c y l i n d r i c a and Fav e l l a s e r r a t a . As t i n t i n n i d s are both p r i m a r i l y herbivorous and c a n n i b a l i s t i c (personal observation); as t h e i r predators i n -clude organisms which are v a r i o u s l y planktonic, benthic, carnivorous or pre-dominantly herbivorous; and as some predators l a t e r become t e r t i a r y carnivores 30 (e.g. f i s h l a r v a e ) , i t can be seen that the microzooplankton may form the base of some ( s t r u c t u r a l l y ) extremely complex food webs. From another point of view, omnivory, which i s probably widespread i n marine plankton,might be con-sidered to have a s i m p l i f y i n g and s t a b i l i z i n g e f f e c t on the dynamics of food webs. Encystment There i s no strong evidence that t i n t i n n i d s or any planktonic marine c i l i a t e s can form r e s i s t a n t and metabolically dormant cysts. This a b i l i t y would probably be useful to t i n t i n n i d s i n t h i s area, but l i k e l y cysts have not been seen during t h i s study. However, s i n g l e thick-walled objects of the appropriate shape have been seen inside the l o r i c a s of F a v e l l a s e r f a t a i n a few samples taken from Georgia S t r a i t and photographed severalyyears a f t e r preservation (F.J.R. Taylor, unpublished data). A few i n d i v i d u a l l o r i c a s of two species of t i n t i n n i d s from the South-West Indian Ocean have been seen to contain plugs j u s t i n s i d e the o r a l end. One of these preserved l o r i c a s also contained what appeared to be a p a i r of r e c e n t l y divided c e l l s (F.J.R* Taylor, unpublished data). Such l o r i c a plugs might be useful to a metabolically quiescent t i n t i n n i d whether the l a t t e r was encysted or not. Loricas with plugs have not been seen during t h i s study. "Photosynthetic" c i l i a t e s U n t i l recently i t has been axiomatic to consider c i l i a t e protozoa as heterotrophs, consuming the complex products of photosynthesis or chemo-synthesis c a r r i e d out by other organisms; or as the hosts of symbionts which are e n t i r e a l g a l c e l l s . During t h i s study, and as a d i r e c t r e s u l t of examin-ing large numbers of f i e l d samples for t i n t i n n i d s , several planktonic c i l i a t e s were found to contain chloroplasts and other 'foreign' organellesqof. a l g a l o r i g i n which were not i n s i d e food vacuoles but 'free' and undigested i n the c i l i a t e cytoplasm. Descriptions of the u l t r a s t r u c t u r e of these organisms and a discussion of t h e i r possible e c o l o g i c a l s i g n i f i c a n c e may be found i n Taylor et ,al_. (1971) and Blackbourn e t . a l . (1973) . Several of these c i l i a t e s also at times ingest and digest whole a l g a l c e l l s , so i f the undigested chloro-p l a s t s prove to be f u n c t i o n a l , the host c i l i a t e s can be considered as both heterotrophs and as f u n c t i o n a l autotrophs. It i s i n t e r e s t i n g and i n e x p l i c a b l e that thesesundirgested chflioroplasts were never found i n s i d e t i n t i n n i d s i n t h i s study, even i n those species with transparent l o r i c a s which might allow the passage of enough l i g h t for photosynthesis. This possible semi-autotrophy amongst some members of the microzooplankton, makes a consideration of marine food-webs even more f a s c i n a t i n g and complex. 32 3) MATERIALS AND METHODS a) Sampling and i n i t i a l treatment A l l t i n t i n n i d s and other microzooplankton used i n experiments were ob-tained by sampling inshore surface seawater from docks or j e t t i e s as close to the time of high-tide as pos s i b l e . Concurrently, temperature and s a l i n i t y observations were recorded and samples were taken for examination and preser-vation. An unconcentrated bucket sample and a short haul with a 30 cm d i a -meter net made of monofilament nylon .mesh of 50 /im aperature were taken. The net haul was made by walking along the dock or j e t t y as slowly as possible (^0.5 m.p.h.) and towing the net j u s t submerged. The net was removed from the water as gently as possible with the cod-end i n a bucket of seawater, and the contents poured gently into a 4 - l i t r e insulated container which had previously been h a l f - f i l l e d with surface seawater. More unconcentrated sea-water was then added to a f i n a l quantity of 3 l i t r e s . If i t was apparent that many zooplankton larger than 400 um had been getted, the p a r t i a l l y d i l -uted net sample was poured gently through a short perspex tube f i t t e d at one end with nylon mesh of 75 yum aperture. The f i l t r a t e from t h i s process was d i l u t e d and placed i n a separate insulated container. About 3 l i t r e s of un-concentrated seawater was placed i n a t h i r d insulated container. The t r i p from dock or j e t t y to the laboratory took from between 2 minutes and 1 hour, depending on l o c a t i o n . The temperature of the water, and the condition of the organisms i n the containers usually remained unchanged for at l e a s t 6 hours. Surface seawater temperatures were measured with an unprotected thermometer, and s a l i n i t i e s were estimated from the s p e c i f i c gravity as meas-ured with a hydrometer. The l a t t e r was checked against an Auto-Lab inductive salinometer and found to d i f f e r from i t by no more than l%oOver the temperature 33 range of the samples. This amount of error i s unimportant given the great tolerance of t i n t i n n i d s to changes i n s a l i n i t y . Transfers of t i n t i n n i d s , other protozoa and c i l i a t e d microzooplankton were made with disposable glass Pasteur pipettes drawn out to a terminal diameter of about 150 yum and f i t t e d with a small hard rubber bulb. Copepod adults and-nauplii. were transferred with pipettes with a mouth diameter of approximately 5yum. Approximately 100 mis of most of the unconcentrated and the netted f i e l d samples were preserved with Lugol's iodine s o l u t i o n . Four drops of the con-centrated s o l u t i o n were added to screw-top glass j a r s , before the sample was added. This ensured the rapid f i x a t i o n of microorganisms i n the-sample. More preservative was added i f the organisms were so numerous as to absorb the i n i t i a l amount of preservative. Preserved samples kept a i r - t i g h t and dark f o r four years showed l i t t l e i n d i c a t i o n of c e l l l y s i s or l o s s of s t a i n . In some experiments where circumstances permitted, formaldehyde buffered with phosphate and d i l u t e d with seawater to a f i n a l concentration of 3%, was used to preserve t i n t i n n i d s . Of the f i x a t i v e s best suited for c y t o l o g i c a l work: gluteraldehyde buffered with phosphate proved unsatisfactory i n t h i s work as a f i x a t i v e and as a preservative, and osmium tetroxide was found to be too dangerous to use i n the working area. The s i z e 'spectrum' of p a r t i c l e biomass (volume) of many f i e l d samples was measured with a Model B Coulter Counter within 1 or 2 hours a f t e r the .(sampQ-re had been c o l l e c t e d (see also Section 3b (iv) ). The t o t a l numbers and volumes of p a r t i c l e s ( l i v i n g and non-living) were estimated f o r several a r -b i t r a r y s i z e classes from approximately 2 yum to 30 /am diameter (Sheldon and Parsons, 1967). 34 The food contents of l i v e t i n t i n n i d s immobilized by c o v e r s l i p pressure were estimated within one or two hours a f t e r c o l l e c t i o n of the sample. More d e t a i l s of these methods are given i n Section 3b ( i i ) . Estimates were made of the number, degree of 'clumping', type, colour, s i z e and state of digestion of food items. The type of food item i n s i d e the t i n t i n n i d s could be e a s i l y i d e n t i f i e d only i n the case of the Bacillariophyceae, Dinophyceae, Eugleno-phyceae, and a small number of other organisms inoother f a m i l i e s of algae. Detritus and protozoans (e.g. other t i n t i n n i d s ) could also be c l e a r l y iden-t i f i e d as ingested food. Approximate estimates were made of the number of i d e n t i f i a b l e s p h e r i c a l food storage granules i n s i d e t i n t i n n i d s ( i . e . 'many', 'few' or 'none') and the approximate average s i z e of suck granules. The l a t -ter estimates could be considered as analogous to measurements of 'condition f a c t o r ' i n la r g e r animals. Since c i l i a t e s are not known to accumulate i n -soluble protein or carbohydrate, these granules may well be composed l a r g e l y of l i p i d s . The t o t a l amount of l i p i d per c e l l , and the number of c e l l s con-ta i n i n g neutral f a t granules i s greatest i n r a p i d l y growing populations of Tetrahymena pyr-Lformis ( H i l l , 1972). L i p i d s are also the major substrates for aerobic metabolism i n t h i s species ( H i l l , l o c . c i t . ) . The presence and s i z e of any early signs of c e l l d i v i s i o n (oral anlage) i n the c i l i a t e s were noted; together with the sizes of the c e l l s and the length of t h e i r o r a l c i l i a . L a s t l y , the dimensions of the t i n t i n n i d l o r i c a s ( i f any) were meas-ured, and the length noted of any portions of any obviously d i f f e r e n t mater-r i a l , which might i n d i c a t e recent l o r i c a a ccretion. The approximate f u l l n e s s of the gut of r o t i f e r s , the pigmentation of t h e i r food and the number of eggs (sexual and asexual) were estimated. Whenever possibl e , the food items were noted f o r at l e a s t 10 of a l l species of t i n t i n n i d s i n a sample. For reasons of s c a r c i t y of for lack of a v a i l a b l e time t h i s was often impossible, 35 and then only 2 or 3 of the les s abundant t i n t i n n i d species were examined. b) Experimental Methods (i ) General comments The feeding rates of organisms can be measured d i r e c t l y or i n d i r e c t l y . D i r e c t methods include ( i ) observation and ( i i ) counts of the number of food items accumulated i n s i d e an organism i n a period of time too short f o r appre-c i a b l e digestion to have feegun. Indirect methods include ( i i i ) the uptake of 14 tracers (e.g. C-labelled algae) and (iv) counts of the number of food items i n the surrounding medium (e.g. with a Coulter Counter) before and a f t e r feeding. Method ( i i ) can only be used i n those organisms such as t i n t i n n i d s which eat r e l a t i v e l y slowly and do not masticate the food before digestion. Direct methods are very accurate and very s e n s i t i v e , but must often be car-r i e d out under r e l a t i v e l y a r t i f i c i a l conditions and can lead to great v a r i -a b i l i t y of r e s u l t s . The best use of i n d i r e c t methods requires considerable knowledge of the physiology of the organisms used; and necessitates, i n the case of slow feeders, the use of r e l a t i v e l y long experiments. However, i n -d i r e c t methods can be used under more natural conditions than some d i r e c t methods and usually give l e s s v a r i a b l e r e s u l t s . A l l four of the above experimental methods were used i n t h i s study: i d i r e c t observation; the counting of accumulated food items; the counting of 14 C-labelled a l g a l food; and Coulter counts of the p a r t i c l e s i n the medium. 14 None was wholly s a t i s f a c t o r y for a v a r i e t y of reasons. The C<~tracer method i n p a r t i c u l a r proved to be too d i f f i c u l t to use w i t h . t i n t i n n i d s due to low rates of ingestion and t h i s technique was soon discarded. The data obtained from d i r e c t observation was sparse and d i f f i c u l t to r e l a t e to other data (see Section 4b). Therefore most of the useful data has been obtained from 36 counts of accumulated food (Section 4a) and from Coulter Counter experi-ments (Section 4c). The experimental organisms (mainly t i n t i n n i d s ) were fed with e i t h e r : n a t u r a l phytoplankton and other p a r t i c l e s from unconcentrated seawater; with one or more laboratory phytoplankton cultures of si n g l e or mixed species composition; or with natural and synthetic i n e r t p a r t i c l e s such as tree p o l l e n and polystyrene l a t e x spheres. Many of the phytoplankton cultures were of l o c a l o r i g i n . Some consistency i n the q u a l i t y of the laboratory phytoplankton food was achieved by using only one type of culture medium, and by only taking sub-samples f or experimental use from cultures which were l e s s than two weeks old and i n the exponential growth phase. This p r a c t i c e should have ensured that the phytoplankton were of r e l a t i v e l y high food value, with r e -l a t i v e l y l a rger amounts of pro t e i n and vitamins etc., than i f the samples had been taken from older cultures. The culture medium used (Jowett's) was based onl'l>oealsseawa'6erpplus a very small amount of s o i l extract and a good balance of n u t r i e n t s . Its composition was as follows: -IN I LITRE 875 mis F i l t e r e d seawater 175 mis D i s t i l l e d water 0.25 G T r i s (Hydroxymethyl)Aminomethane 0.10 G NaNO 3 0.01 G K 2HP0 4 Na„Si0 o+10 mis IN HCL 0.03 G 6 x 10 3G 2 x 10 G FeC7l_6Ho0 37 2 x -4 10 G Mn(as SO^) 2 x 10 _ 5G Zn(as Cl) 2.5 x 10~ 6G Co(as Cl) 2.5 x 10~ 6G Cu(as Cl) 3.9 x 10~ 4G Mo (as Na) 3.0 x 10" 4G H 3 B o 3 5 x -4 10 G Thiamine HCL 3 x 10" 5G N i c o t i n i c Acid 3 x 10~ 5G Ca Pantothenate 3 x 10" 6G P-Aminobenzoic Acid 1 X 10 _ 6G B i o t i n 5 x -4 10 G I n o s i t o l 6 x 10~ 7G F o l i c Acid 1 X 10" 6G Cyanocobalomin 2.5 mis S o i l Extract (Supernatant from autoclaved mixture of equal volumes of s o i l and d i s t i l l e d water) The experimental containers were normally p l a s t i c or glass beakers of 50 to 500 ml volume, and these were usuallyccovered with aluminum f o i l to exclude the l i g h t . The containers were rinsed and scrubbed i n d i s t i l l e d water only. A few experiments were done with organisms i n o n e ^ l i t r e p l a s t i c stoppered b o t t l e s t i e d to the side of f l o a t i n g j e t t y i n Saanich i n l e t . Most experimental containers were placed i n trays i n incubators held f o r other reasons at 9 C, 13 C or 16 C or at room temperature (about 22 C). Whenever pos s i b l e , the containers were placed on a slowly-reciprocating shaker t a b l e . Experiments la s t e d from a few minutes to several hours. Food p a r t i c l e s of laboratory o r i g i n were dispensed into the experimental containers from s i n g l e -species stock cultures with Eppendorf micropipettes or small volume glass 38 pipettes. The numbers and average volumes of the c e l l s i n these stock c u l -tures were measured with the Coulter Counter from 1/100 d i l u t i o n s immediately before the experiments. In many of the experiments, t i n t i n n i d s were gently pipetted onto glass s l i d e s f o r the examination of t h e i r food contents with a high-power microscope at i n t e r v a l s or at the end of the experiment. As these t i n t i n n i d s were thereby k i l l e d , i t was impossible to make repeated ob-servations on the same organisms and thus each sample was taken as represent-ati v e of the experimental population at that time. On occasions, sub-samples of t i n t i n n i d s taken at i n t e r v a l s were fi x e d and preserved f o r 1 or 2 days i n formaldehyde before examination, but t h i s was l i m i t e d to a minority of experi-ments invo l v i n g food organisms that were e a s i l y d i s t i n g u i s h a b l e when preserved. ( i i ) Counts of accumulated food The food contents of l i v e microzooplankton were estimated from c e l l s immobilized by c o v e r s l i p pressure i n small volumes of water under t h i n cover s l i p s on glass s l i d e s . Eventually c i l i a r y beating i n these c e l l s ceased, and c e l l l y s i s began. Larger organisms such as r o t i f e r s were often squashed by t h i s procedure, and t i n t i n n i d s were often extruded from t h e i r l o r i c a s . The c e l l plasma membrane of extruded c e l l s did not n e c e s s a r i l y break, nor did c e l l l y s i s occur immediately i n squashed organisms. The c e l l contents of t i n t i n n i d s with non-transparent l o r i c a s often could not be seen c l e a r l y unless some extrusion of the c e l l from the l o r i c a had taken place. Also, protozoa containing many food items could only be c l e a r l y examined a f t e r some extrusion and f l a t t e n i n g had occurred. The estimation of the proportion of ingested food items which had undergone d i g e s t i o n was somewhat subjective under these circumstances. However, a f t e r considerable experience i t became c l e a r that undigested food items did not change r a p i d l y i n appearance u n t i l a f t e r t i n -t i n n i d c e l l l y s i s began, and the l a t t e r followed by several seconds the 39 the complete cessation of motion of the t i n t i n n i d adbral c i l i a . Therefore, 'semi-digested' c e l l s were considered to be those appearing d i s t o r t e d i n shape, changed i n colour, reduced i n s i z e , etc., before the c i l i a r y organelles stopped beating. A p a r t i a l l y digested food item can be confused with an un-digested item of smaller s i z e and d i f f e r e n t shape, and d i s c r i m i n a t i o n between the two was e n t i r e l y subjective and based on previous experience. This problem was n a t u r a l l y most d i f f i c u l t when natural food items were examined, p a r t i c u l a r l y when they were d i f f i c u l t to r e l a t e to i d e n t i f i a b l e items i n the environment. The c e l l structures most r e s i s t a n t to the d i g e s t i v e enzymes i n the food vacuoles of t i n t i n n i d s included the s i l i c e o u s c e l l walls of diatoms, the c e l l u l o s e thecae of d i n o f l a g e l l a t e s , and the c a r o t e n o i d - l i p i d 'eyespots' of some f l a g e l l a t e s . These are probably a l l egested eventually having under-gone l i t t l e d i g e s tion. Of these structures, only the reddish (and often clum-ped) eyespots could be sometimes confused with other small whole food items. I t i s i n t e r e s t i n g that one (but apparently only one) l o c a l c i l i a t e , a species of the O l i g o t r i c h genus Strombidium, seems to r e t a i n these a l g a l eyespots for i t s own use (Blackbourn e t . a l . , 1973). These clumped eyespots might i n some future work prove to be useful long term markers of the rates of ingestion and d i g e s t i o n of c e r t a i n food items. This would be best c a r r i e d out i n con-j u n c t i o n with observations with the electron micE©'scope, which was impossible to use i n t h i s study. ( i i i ) Observations of feeding behaviour Most observations were made with a Mark I I I Zeiss Stereomicroscope f i t t e d with a zoom focussing control and sub-stage i l l u m i n a t i o n . The range of t o t a l magnification was x4 to x40. The t i n t i n n i d s swam f r e e l y i n shallow P e t r i 40 dishes placed on the microscope stage and containing 10 to 15 ml of seawater to a depth of 0.7 to 1.0 cm. The transmitted double l i g h t source was c a l i -brated with a Lightmeter (Photovolt Corp.) equipped with neutral f i l t e r no. 3. For most observations the l i g h t i n t e n s i t y at the microscope stage was approximately 2,000 Lux. The l i g h t passed through a heat f i l t e r but seawater i n the P e t r i d i s h would warm from about 10 C to about 20 C i n about 30 min-utes. Therefore, t i n t i n n i d s and/or seawater were taken from insulated con-tainers for short observation periods only. T i n t i n n i d s showed l i t t l e apparent r e a c t i o n to the i l l u m i n a t i o n , and at a l l l l i g h t i n t e n s i t i e s tended to move upwards to the water surface unless under obvious p h y s i o l o g i c a l s t r e s s . There i s l i t t l e or no d i f f e r e n c e i n the food accumulated by t i n t i n n i d s i n strong, dim or no l i g h t . This has been shown i n experiments (see Section 4a); and i n samples from Coal Harbour where the food contents of t i n t i n n i d s taken at dusk and again at dawn, showed e s s e n t i a l l y no d i f f e r e n c e s . Goulder (1973) also found no d i u r n a l p e r i o d i c i t y of feeding i n the freshwater planktonic c i l i a t e s Loxodes s t r i a t u s and Loxodes  magnus. Therefore, i t i s assumed that the i n t e n s i t y of l i g h t had no e f f e c t on t i n t i n n i d feeding behaviour during these observations (but see Results i n Section 4b). The zoom rapid focussing control was e s s e n t i a l f or c l o s e l y following a p a r t i c u l a r i n d i v i d u a l t i n t i n n i d , as i t changed d i r e c t i o n frequently and r a p i d l y and moved i n shallow h e l i c e s . The r i g h t hand was used to c o n t r o l the focus and the p o s i t i o n of the P e t r i d i s h on the microscope stage, and the l e f t hand c o n t r o l l e d 2 stopwatches and a counting device. Events were recorded as follows: the t o t a l length of observation of a p a r t i c u l a r t i n -t i n n i d (a 'run') was timed with stopwatch a ) , and the duration of one event of i n t e r e s t (e.g. the 'handling' of a food item by the t i n t i n n i d ) during a 'run'.was timed with stopwatch b). The counter was used to record a) the number of apparent p a r t i c l e s contacted by a t i n t i n n i d per run, and b<) the number of such contacts which res u l t e d i n ingestion by the t i n t i n n i d . The smallest p a r t i c l e that could be detected and i t s f a t e c l e a r l y followed, was of approximately 4/m diameter (but see Section 4a and 4 b). The stopwatch and counter records were noted at the end of each run and they were reset. When possible, 5 to 10 i n d i v i d u a l s of each species were observed one at a time u n t i l t h e i r behaviour changed d r a s t i c a l l y , e.g. by staying at the bottom or at the water surface of the dish. Occasionally, a f t e r a 'run', the t i n t i n n i d was placed on a s l i d e f o r examination of the food contents at high magnification. Where a p a r t i c l e was large enough to be seen during ingestion; or large enough to cause the t i n -t i n n i d to stop moving or to change d i r e c t i o n i n order to handle i t , whether by ingestion or eventual r e j e c t i o n , the event could be accurately recorded. However, some ing e s t i o n may have gone unrecorded. Very small or unfocussed items might well have been ingested with no overt behavioural change by the t i n t i n n i d ; and i n the opposite d i r e c t i o n , minor changes i n the angle of the path of t i n t i n n i d movement may not always have indicated ingestion or r e j e c t -ion. (iv) Coulter Counter experiments The method used i s e s s e n t i a l l y s i m i l a r to that described by Sheldon and Parsons (1967). A Model B e l e c t r o n i c Coulter Counter was used with aperture tubes of 50 /im or 100yum diameter o r i f i c e s . The numbers and t o t a l volumes were calculated f or p a r t i c l e s of between 1.2 and 20.0 yum diameter. With t h i s technique, the numbers of p a r t i c l e s i n each of a continuous seri e s of a r b i t r a r y 42 s i z e classes expanding geometrically, were m u l t i p l i e d by the average volumes (calculated as spheres) of each s i z e c l a s s . Hence, the diameters corres-ponding to the average spheres i n these s i z e classes were as follows: 1.78, 2.24, 2.82, 3.57, 4.49, 5.66, 7.12, 8.98, 11.3, 14.3, and 18.6^um. This t o t a l 'spectrum' covered the s i z e range of most p a r t i c l e s eaten by most t i n t i n n i d species i n t h i s study, but of course most natural p a r t i c l e s are non-spherical. The use of a s p h e r i c a l average volume was unavoidable, but i t should be noted that the actual dimensions of a food p a r t i c l e would be of importance to a t i n t i n n i d , p a r t i c u l a r l y near the upper s i z e l i m i t . o f i t s feeding a b i l i t y . Counts were made on subsamples taken at the beginning and end of the experiments from both 'experimental' and 'control' containers and were de-signated as E^, E^, C^ and C^ r e s p e c t i v e l y . Control containers were set up from f i e l d samples by passing 200 to 500 ml of the sample through nylon mesh of a s i z e (usually 30 um for t i n t i n n i d s ) to exclude a l l predators, and i n -c i d e n t a l l y also much large phytoplankton, usually diatoms. U n f i l t e r e d water, or water f i l t e r e d through mesh of a s i z e to exclude only predators larger than t i n t i n n i d s , was used i n the 'experimental' containers. The subsamples were slowly s t i r r e d during counting which took from 10 to 30 minutes to com-ple t e . Six r e p l i c a t e counts were taken i n each s i z e range, with a variance of 5 to 10% of the mean value for counts between 100 and 1000, and a variance of 10 to 20% of the mean for counts between 10 and 100. Counts were taken for 2 to 16 seconds, depending on the frequency of p a r t i c l e s . The accuracy of the estimation of p a r t i c l e volume by the counter was checked on a number of occasions by c a l i b r a t i o n with ragweed pol l e n of known s i z e . The accuracy of the estimation of p a r t i c l e number was not checked, as a l l other counting methods were considered to be l e s s accurate, and so has 43 been assumed to be absolute. For t h i s reason perhaps, the Coulter Counter i s best used for the estimation of r e l a t i v e changes i n number, as i n these experiments, rather than for counting the numbers of p a r t i c l e s i n a f i e l d sample. After the f i n a l Coulter count, the predators i n the subsample from the experimental container were counted. The dead organisms were counted at x20 magnification on a squared P e t r i dish before the subsample was f i x e d . Immediately a f t e r f i x a t i o n , a l l the organisms were counted, and the d i f f e r e n c e between the two counts was considered to be the l i v e r t o t a l . This was done for three 10 ml subsamples of the container, and the mean value c a l c u l a t e d . The volumes i n each s i z e c l a s s and i n the t o t a l , were calculated for C^, C^, E^, and with the use of a computer programme. This programme also tested the hypothesis of no d i f f e r e n c e between i n i t i a l and f i n a l values i n each s i z e c l a s s <H= 0.05). As the frequency of occurrence of a p a r t i c l e i n seawater often decreases r a p i d l y with increasing s i z e , the counts i n the largest s i z e classes were r e l a t i v e l y low and the values calculated were often not s i g n i f i c a n t as a r e s u l t . To count any s i z e c l a s s for more than 16 seconds would have been unduly time-consuming, and longer counts were not made. This d e c i s i o n may have resulted i n the underestimation of feeding rates on the l a r g e s t s i z e classes i n some experiments. The t o t a l p a r t i c l e volumes for C^ and never d i f f e r e d by more than 20%. Two values for the feeding rate were estimated for reasons described below, but both c a l c u l a t i o n s were made with an equation derived from those given by Frost (1972) and others. A combination of 3 equations i n Frost (1972) (as modified by my notation) may be written as follows: FR/P = E ( : e ( k _ g ) r - l ) Vg — — C D T.(k-g).P 44 where FR/P = ingestion r a t e as c e l l s eaten/predator/hour P = number of predators^ i n experimental container V = volume of experimental container k = a l g a l growth c o e f f i c i e n t i n control container g = grazing c o e f f i c i e n t i n exper-imentalacontainer and T = duration of the experiment i n hours. Equation (1) can be written as: (2) FR/P = ((E 2-E 1)/;(T.P.)). (log 10 ( C 2 / C 1 ) - l o g 10 ( E ^ E ^ / l o g 10 ( E ^ ) ) 3 where FR/P = volume (jam ) of p a r t i c l e s eaten/predator/hour. and P = number of predators/ml i n experimental container. It w i l l be seen that equation (2) assumes that the grazing mortality i n E, and the increase i n numbers ( i f any) of the food organisms i n both C and E i s exponential. If these changes were l i n e a r i n form the corresponding equation would be written as: FR/P = ( ( ( C 2 . E 1 ) / C 1 ) - E 2 ) / T . P (3) These two equation can give very d i f f e r e n t values for FR/P. The use of equa-t i o n (2) w i l l give lower values of FR/P than equation (3) when there i s any (exponential) net increase i n the control population during the experiment; equal values of FR/P when there i s no net increase or decline i n the c o n t r o l ; and higher values of FR/P than equation (3) when there i s a net decline i n both the c o n t r o l and experimental containers. Id e a l l y , the form of the grazing mortality c o e f f i c i e n t and the growth c o e f f i c i e n t , should be estimated from samples taken from both C and E at frequent i n t e r v a l s throughout an experiment (see Frost, 1972). This i s not 45 f e a s i b l e with slowly feeding organisms such as t i n t i n n i d s . A l t e r n a t i v e l y , there should e x i s t some independent estimate of the maximum feeding rate of the organism under conditions s i m i l a r to those pertaining i n the experiment. Unfortunately, there was l i t t l e r e l i a b l e data of t h i s type a v a i l a b l e i n t h i s study (see Section 4b). The p a r t i c l e concentration (number or volume/ml) at which an organism reaches i t s constant maximum ingestion r a t e at a given temperature and for a p a r t i c u l a r type of food, i s termed i n t h i s study the 'optimal food concentration' (OFC). This change i n rate may be abrupt or gradual. The correct choice of the method of c a l c u l a t i o n of feeding rates r e l i e s upon a knowledge of the OFC. Most invertebrate f i l t e r - f e e d e r s are generally assumed ( u n t i l behavioural or s e l e c t i o n experiments are done) to sample t h e i r environment passively and completely, under 'normal' circumstances. That i s , contact i s made with food items and they are a l l eaten, at a c h a r a c t e r i s t i c constant rate at a given temperature. This leads to an exponential rate of decline of a non-growing population of food c e l l s , and equation (2) may then be used to c a l c u l a t e f f e e d i n g r a t e s . At any food concentration above the OFC bel'ow some i n h i b i t o r y l e v e l , the organism w i l l feed at i t s maximum rate no matter what the concentration, and the l a t t e r w i l l d e cline slowly and l i n e a r l y . In such circumstances, equation (3) may be used to c a l c u l a t e feeding r a t e s . If the population of grazed food items decreases from a concentration above the OFC to a concentration below the OFC during the experiment, then a l i n e a r rate of decline w i l l be followed by an exponential r a t e of de c l i n e . The form of the natural rate of increase i n number of a growing pop-u l a t i o n of c e l l s can be l i n e a r , exponential or hyperbolic, depending upon the environmental conditions and i n t e r n a l p h y s i o l o g i c a l state of the c e l l s . When 46 c e l l growth i s rapi d , the exponential form of population increase i s most l i k e l y , and has been assumed to be true i n a l l these experiments. Hence the use of equation (2). Some of the r e s u l t s i n d i c a t e a_ p o s t e r i o r i that the OFC had not been exceeded i n these Coulter Counter experiments, and that therefore the choice of equation (2) had been corre c t (but see Section 4a). If any s i z e and/or taxonomic group of phytoplankton reproduced so r a p i d l y during the experiment so as to push the t o t a l p a r t i c l e volume i n the experimental vessel temporarily over the OFC l e v e l f o r a t i n t i n n i d species, the l a t t e r may have al t e r e d i t s feeding behaviour i n response, thus increasing the l i k e l i h o o d of poor c o r r e l a t i o n s between the calculated feeding rate and the logmean experimental t o t a l food volume (see Section 4a and General Discussion). The value of the t o t a l feeding rate (FR/P) for each experiment was i n i -t i a l l y c a l c u l ated as the sum of negative and p o s i t i v e FR/P values i n each s i z e c l a s s . This c a l c u l a t i o n , 'net t o t a l consumption' (or NTC), gave anomalous p o s i t i v e values i n several experiments. That i s , i n those experiments the predators appeared to make a net a d d i t i o n of p a r t i c l e s to the medium r e l a t i v e to the c o n t r o l . Poulet (1974) also found p o s i t i v e values i n some experiments using the Coulter Counter. There are a number of possible explanations for such r e s u l t s which include (1) contamination of the experimental container (only) with extraneous p a r t i c l e s during the experiment; (2) the e f f e c t of pre-dators on a) resuspending sedimented p a r t i c l e s ; b) increasing the growth of algae i n some s i z e classes by the excretion of NH^, etc.; c) increasing the growth of algae i n some s i z e classes by s e l e c t i v e l y removing other algae wMch are competing with them for scarce n u t r i e n t s ; or d) creating l a r g e r or smaller p a r t i c l e s by clumping or comminution from those p a r t i c l e s that they attempt to ingest. Most of the Coulter Counter experiments were too lengthy (20 to 52 hours) to r u l e out any of" these p o s s i b i l i t i e s , although (2) b) and (2) c) seem the most l i k e l y . In many experiments r e l a t i v e p a r t i c l e removal (consumption) was demonstrated i n a l l or most s i z e c l a s s e s , and i n the r e -mainder no net change i n t o t a l volume was seen. The p o s s i b i l i t i e s mentioned above may apply even i n experiments where net t o t a l consumption occurred. Also, a p a r t i c u l a r s i z e class may have had better growth or l e s s m o r t a l i t y by chance i n the experimental container than i n the con t r o l container. Food s e l e c t i o n experiments (see Section 4a) have indicated that some items may not be eaten at a l l by some t i n t i n n i d s , and that other items may be eaten to varying degrees. This may be r e f l e c t e d i n those s i z e classes which show no r e l a t i v e change between the control and experimental containers. It must be remembered that the s i z e classes used are a r b i t r a r y , and that natural or laboratory food items of one species may spread over several Coulter s i z e classes. One s i z e c l a s s may also contain several d i f f e r e n t species of food item, e s p e c i a l l y i n natural samples. A l l i n a l l , the calculated (NTC) values of FR/P may represent minimal net values of consumption by t i n t i n n i d s , but reasonably r e f l e c t t h e i r o v e r a l l t o t a l e f f e c t s on natural phytoplankton pop-u l a t i o n s . This may be true even i n those experiments where a l l s i z e classes show net consumption, i f some food items i n those s i z e classes are at the same time unaffected by t i n t i n n i d feeding. To obtain values for FR/P close to the maximum possible f o r a l l experiments i t was also calculated as the t o t a l of a l l negative (consumed) values only, and these values were desig-nated as 'edible spectrum o n l y 1 (or ESO). Mult i p l e c o r r e l a t i o n c o e f f i c i e n t s were calculated separately for NTC and ESO values, f o r 10 variables a s s o c i -ated with these experiments (see Section 4c). One of the var i a b l e s expected to be strongly correlated with values of FR/P was C2^C1' o r t h e 8 r o w t l> rate of the control population, which i s assumed 48 to be also the p o t e n t i a l growth rate of the populations of food c e l l s i n the experimental container. This assumption i s l e a s t j u s t i f i e d i n the longest experiments. Also, ^2^1 values are the mean of the t o t a l spectrum, and these would not ne c e s s a r i l y c o r r e l a t e well with t o t a l FR/P values calculated over the whole spectrum. As a check on the p o s s i b i l i t y that the 'feeding size-spectrum' of t i n t i n n i d s i s enlarged when l e s s t o t a l food i s a v a i l a b l e , i . e . that they become l e s s ' s e l e c t i v e ' when 'hungry', the number of Coulter s i z e classes showing net consumption f o r each experiment (ESP) was included as a v a r i a b l e (4) i n the c a l c u l a t i o n of c o r r e l a t i o n . Variable (3) i n Table 27 i s 'logmean E' - which i s considered to be the mean value of t o t a l t f o o d a v a i l -able to the predator during the experiment, adjusted to an experimental dur-at i o n of 24 hours. This value i s transient and for long experiments with large changes i n C and E p a r t i c l e concentrations i t i s also a r b i t r a r y . It was calculated as follows: E2 " E l logmean E = (4) ^ 2 " L o & n E l The use of th i s equation gives very s i m i l a r r e s u l t s to that of equation (3) i n Frost (1972) . GLOSSARY 49 the t o t a l a i n i t i a l and f i n a l p a r t i c u l a t e volumes from 1 to 20 /im i n the con t r o l v e s s e l . as above for the experimental v e s s e l . 3 i n d i v i d u a l feeding rate i n numbers of c e l l s or^um /pre-dator /hr or as equivalent ml/pred/hr. optimum food concentration - the food concentration at which the feeding rate of a predator i s a t , or close to, maximum. net t o t a l consumption - the t o t a l feeding rate (FR/P) from a l l s i z e classes measured, including net additions or losses of p a r t i c l e volume. edi b l e spectrum only - the t o t a l feeding rate (FR/P) from only those s i z e classes showing net losses of p a r t i c l e volume (Consumption). index of increase of t o t a l p a r t i c l e volume i n the con t r o l v e s s e l during the experiment. edible spectrum portion - the number of s i z e classes i n an experiment which showed net losses of p a r t i c l e volume. logarithmic mean value of p a r t i c l e volume a v a i l a b l e to the predator. An index of p r o p o r t i o n a l i t y of ingestion of a p a r t i c u l a r item i n the d i e t compared with i t s abundance i n the medium. the dif f e r e n c e between the e l e c t i v i t y indices of various prey types with no comparison with a single-prey s i t u a t i o n . No s e l e c t i o n or preference i s i n f e r r e d . 50 Negative Sel e c t i o n Loss Rates Search Rate Contact Rate (CR) Control of P Phytoplankton G rowth where a food type has a neutral e l e c t i v i t y index i n one s i t -uation and a strongly negative index i n another s i t u a t i o n where more prey types are involved. However, where a l l prey types show a more negative index when presented together than when presented with fewer other prey types, then the cause may l i e i n the f a c t that the t o t a l prey concentration i s now above the OFC for that predator. In long experiments apparent feeding s e l e c t i o n may be the r e s u l t of differences i n prey d i g e s t i b i l i t y . the rate of apparent disappearance of food from t i n t i n n i d s 3 i n yum /pred/hr. the t h e o r e c t i c a l rate at which the medium i s thoroughly searched by a predator, i n ml/pred/hr. the rate at which a p a r t i c u l a r concentration of prey i s contacted by a predator i n nos/pred/hr or equivalent ml/pred/hr. E2 C2 when -=—<l and — > 1 o v e r a l l , as the net r e s u l t of r e l a t i v e E 1 C 1 t o t a l p a r t i c l e volume changes i n 24 hours. 51 4) RESULTS AND DISCUSSION a) Accumulation Experiments ( i ) Q u a l i t a t i v e Results Table 1 presents some of the c e l l measurements of the t i n t i n n i d species studied; and from t h i s table the r e l a t i o n s h i p between the volume of the l a r g -est s i n g l e contained food item, whether i d e n t i f i e d or not, and the maximum estimated volume of the t i n t i n n i d c e l l s i s shown i n Figure 5. In almost a l l cases, the maximum volume of t i n t i n n i d c e l l s r e f e r s to the maximum s i z e of the parental c e l l j u s t before d i v i s i o n . The minimum volume of the r e s u l t i n g daugh^r c e l l s w i l l be about h a l f the volume of the parental c e l l . A two-fold d i f f e r e n c e i n the c e l l volume of a species w i l l a l t e r Figure 5 very l i t t l e . In Figure 5 the r e l a t i o n s h i p i s described by the equation: log t i n t i n n i d 3 3 volume ( i n yum ) = 2.392 + 0.757 l o g food volume (i n jsm ). The s i g n i f i c a n c e of t h i s g e n e r a l i t y i s unknown. The species whose value f a l l s furthest from the l i n e i s Tintinnidium mucicola. Other unusual features of the feeding be-haviour of t h i s species w i l l be discussed l a t e r . There appears to be no absolute minimum food s i z e f o r t i n t i n n i d s (see Table 2) unlike other zoo-plankton (Parsons, et^al.^, 1967; Poulet, 1973); but very small food items may be eaten i n equal or greater proportion to t h e i r abundance much l e s s consis-t e n t l y than larger items, by at l e a s t one species of t i n t i n n i d (see Section 4c). As a f i e l d sample may contain up to ten species of t i n t i n n i d s , the s i z e overlap i n t h e i r feeding may be considerable. Those t i n t i n n i d species 4 3 i n the narrow range of volumes between 3 and 10 x 10 yum show almost no r e l a t i o n s h i p between c e l l volume and volume of largest food item. For example Tintinnopsis subacuta i s l e s s than twice as large as Tintinnidium mucicola, but the d i f f e r e n c e i n the volume of t h e i r l a r g e s t food item i s almost ten-f o l d . The la r g e s t item seen-inside both Tintinnopsis subacuta and CM m T A B L E I. List of Tintinnid species and their cell measurements. Species Lorica Dimensions ( r i m ) Diam. Length 'Ave.' Max Eutintinnus latus 75 200 250 E . tubulosus 25 100 150 Favella serrata 120 250 250 Helicostomella kiliensis 30 150 250 Ptychocyclis acuta 70 100 100 Stenosomella ventricosa 70 90 100 S. nivalis 30 50 50 Tintinnidium mucicola 45 150 350 Tintinnopsis subacuta 45 100 160 T . cylindrica 40 160 220 T . parvula 35 80 150 T . rapa 25 60 75 T . nana 20 40 60 Maximum Cell Dimensions (|im) Diam. Length 70 20 100 25 60 50 25 35 40 35 • 30 20 15 200 100 200 100 80 100 30 60 80 90 70 40 25 Cell Volume 5 x 10 2.5 x 104 8 x 105 4 x 104 8 x 104 8 x 103 5 x 104 7 x 104 7 x 104 3 x 104 6 x 103 3 x 103 Length of Adoral Cilia (Um) 35 20 50 30 50 30 30 (thin) 35 3 5 (thin) 35 10 10 53 Figure 5. Relationship between t i n t i n n i d c e l l volume (TV) and maximum observed volume of i n d i v i d u a l food items (FV). 1 0 ' 10" T V ( P m 3 ) 10 10* 10 E u t i n t i n n u s l a t u s • , • F a v e l l a s e r r a t a L o g T V = 2 . 3 9 2 • 0 . 7 5 7 L o g F V S t e n o s o m e l l a v e n t r i c o s a * T i n t i n n i d i u m 9 e H ^ T I c o s t o m e l l a k i l i e n s i s ^ T i n t i n n o p s i s s u b a c u t a • T . c y l i n d r i c a 'T . p a r v u l a ^ E u t i n t i n n u s t u b u l o s u s r e n o s o m e l l a n i v a l i s " i n t i n n o p s i s r a p a r ° T. n a n a 1 0 ' 1 0 ' F V ( p m J ) 1 0 ' 1 0 < 55 Tintinnopsis c y i i n d r i c a was Tintinnopsis nana, during a 'bloom' of the l a t t e r . These medium-large species are probably the most important of a l l t i n t i n n i d s i n t h e i r grazing e f f e c t s on phytoplankton f o r most of the year. The r e s u l t s of q u a l i t a t i v e and quantitative tests of the feeding a b i l i -t i e s of t i n t i n n i d s on known food items are presented i n Table 2. In t h i s table also the approximate value of the upper food s i z e boundary and the complete lack of a lower food s i z e boundary i s evident. In comparison the 8 3 freshwater c i l i a t e St en tor coeruleus (c*2 x 10 yum ) eats prey ranging i n s i z e from b a c t e r i a to fellow Stentors (D.J. Rapport - personal communication). Table 2 also shows the wide range i n q u a l i t y of the p a r t i c l e s eaten by t i n -t i n n i d s . L i v i n g and i n e r t items are ingested by a l l species for which they are not .too large. Willow (Salix) and Yew (Taxus) p o l l e n , wine yeast c e l l s and corn starch granules are a l l at l e a s t p a r t l y digested by those t i n t i n n i d s which ingest them. Latex spheres are not digested, but are compacted i n large boluses i n the c e l l a f t e r one or two hours. The only p o t e n t i a l food items to which healthy animals show no apparent feeding r e a c t i o n are the colo u r l e s s f l a g e l l a t e s and b a c t e r i a which are present i n large numbers i n crowded net f i e l d samples f u l l of dead and moribund zooplankton. However, t i n t i n n i d s are often seen to v i o l e n t l y r e j e c t p a r t i c l e s which are small enough for them to ingest and which appear innocuous to the observer. Several spec-ies of laboratory phytoplankton are shown i n Table 2 as 'variably' eaten by t i n t i n n i d s , p a r t i c u l a r l y : Pyramimonas c . f . g r o s s i i , Pavlova gyrans, and Brymnesium parvum. In most cases these species were not eaten when used i n 4 experiments i n heavy f i n a l concentrations (>10 cells/ml) or from old (about 1 month) stock cultures. 'Always' and 'never' eaten r e f e r s to at l e a s t three 4 te s t s at moderate « 1 0 /ml) concentrations. Such species as Monochrysis 56 T A B L E 2. Food eaten by microzooplankton. E = always eaten; V = variably eaten; N = never eaten; blank = untested. F O O D P H Y T O P L A N K T O N M I C R O Z O O P L A N K T O N Tintinnids Others S P E C I E S CLASS V o l . (Um 3) id c id c rS a a u H to .—i > s cn CO 3 CQ O 3 _o 3 +j C4 3 > u n a. 01 cn C 0} X X td o V (j 3 £ H n u -a c >. o d 3 U cj -Q 3 cn ni cn o a tc a > cn 3 <d y it ia u u a tn c cr C c c t c a c <r a > tr n I 1 u o 1 -ti j -1 w Pseudocalanus minutua N III & IV Barnacle nauplii Micromonas sp. Unidentified Monochrysis lutheri Isochrysis galbana Coccolithus huxleyi . Thalas siosira pseudonana Isoselmis sp. Plagioselmis sp. Phaeodactylum tricornutum Pyramimonas c f . gross i i Pavlova gyrans Pry nine slum par vum Dunaliella tertiolecta Platymonas maculata Cr yptomonas • minuta Pseud ope dineila pyrifor mis Eutreptiella sp. Amphidinium carterae C r yptomonas profunda Prasinophyceae Blue-green Bacterium Haptophyceae Haptophyceae Haptophyceae Bac i l lar io -phyceae Cryptophyceae Cryptophyceae Baci l lar io-phyceae Prasino • Haptophyceae Haptophyceae Chiorophyceae Chlorophyceae Cryptophyceae Chrysophyceae Euglenophyceae Dinophyceae Cryptophyceae 3 20 50 50 60 60 75 130 130 140 150 150 200 300 450 450 500 800 1200 V V N N N N N N N E V V N N N N N E E V V N V N N E E E V N E V N V N E E E V V V V V V V N V v N E E E E V N E E E v V v E E N N N V N N V N N N N E E E V N E E N V N V N E E E E E E E E V V V V TT E E V E V E E E E E E E V V V V E E E V E E E V E V N E E V V E E V v i E E E E l E; N N N N N E E E N E E V E E E E E V v v E E E E E E O T H E R FOOD Kaolin Polystyrene Polystyrene Polystyrene Polystyrene Salix and Taxus Yeast cells Starch grains Latex Latex Latex Latex Pollen 1-4 0.5 17 380 3000 1-3000 50-3000 30-3000 E V N E E E E V E E E V V E E E V V V !•: 1 E E E V V V E Jv •i !l E ' i N j i E E E E V E N 57 l u t h e r i , Isochrysis galbana, D u n a l i e l l a t e r t i o l e c t a and E u t r e p t i e l l a sp. were eaten i n almost any concentration or condition by most species; and con-sequently the l a t t e r were much used i n the quantitative experiments on feeding and l o s s rates and feeding preferences, described i n Section 4a ( i i ) . T i n - tinnidium mucicola and Eutintinnus latus showed anomalous feeding preferences or a b i l i t i e s f o r t i n t i n n i d s of t h e i r s i z e , (see Table 2) and these a b i l i t i e s have been examined experimentally (see Sections 4a and 4b). In the majority of f i e l d samples Tintinnidium mucicola contained food items of 5 to 10 jam diameter which mostly had the appearance of cryptomonad c e l l s or chloroplasts,. Many of the quantitative accumulation experiments i n Section 4a are concerned with the apparently s e l e c t i v e feeding behaviour of the common group of species of middle s i z e : Tintinnopsis subacuta, T_. c y l i n d r i c a , Stenosomella ventricosa and Tintinnidiumfaimucicola. Whether t i n t i n n i d s and other zooplankton can or do ingest b a c t e r i a and/ or d e t r i t u s i s a question of some importance to an understanding of the sur-v i v a l of zooplankton i n food-poor s i t u a t i o n s ; such as i n some t r o p i c a l seas, below the euphotic zone, or i n winter i n high l a t i t u d e s . Almost a l l of these t i n t i n n i d species including the la r g e s t were seen to contain a few b a c t e r i a and small d e t r i t a l p a r t i c l e s , most usually i n conditions where concentrations of phytoplankton were f a i r l y low. This question requires a great deal more study. Tintinnopsis nana and T. rapa must almost c e r t a i n l y depend upon bac-t e r i a and phytoplankton of les s -ljhan44/im diameter for t h e i r p a r t i c u l a t e nut-r i t i o n , since they appear to eat nothing much larger than t h i s . I t was unusual to f i n d either of these species i n a f i e l d sample to contain any v i s i b l e food items. However, T_. nana was frequently very numerous (>15/ml) i n t h i s study, but even then appeared to contain l i t t l e food. I t i s possible that T_. nana and _T. rapa obtain food from organic material, either dissolved 58 or more l i k e l y absorbed into very small p a r t i c l e s of d e t r i t u s . Useful d i s -solved material may be i n f a i r l y high concentrations i n t h i s area, but since T_. nana r a r e l y contains p a r t i c l e s , and 'contacts' a very small volume of water and number of p a r t i c l e s i n an hour (see Section 4a), t h i s source of n u t r i t i o n i s also dubious. The l o r i c a of Tintinnopsis nana i s normally cov-ered with small p a r t i c l e s of inorganic d e t r i t u s , as are the l o r i c a s of a l l the members of that genus. When T_. nana i s ingested by larger t i n t i n n i d s or other predators much of the d e t r i t u s i n s i d e the l a t t e r can be a t t r i b u t e d to t h i s source, and not to the ingestion of i n d i v i d u a l small p a r t i c l e s . An i n d i c a t i o n of the permeability of the plasmalemma of t i n t i n n i d s to dissolved substances was obtained by adding large amountsoof 0(a)nneutralrred and (b) neutral red and methylene blue dyes to a f i e l d sample. Two hours l a t e r the t i n t i n n i d s Tintinnopsis subacuta, Stenosomella ventricosa and T i n - tinnidium mucicola were seen to have dye only i n s i d e food vacuoles containing food items. The food items and possibly also the soluble contents of the food vacuoles, were stained. The degree of s t a i n i n g of the O l i g o t r i c h c i l i -ates i n the sample was uncertain. D i n o f l a g e l l a t e c e l l s were generally but f a i n t l y stained, and R o t i f e r s showed st a i n i n g only i n the e p i t h e l i a l c e l l s . Copepod adults and n a u p l i i and gastropod larvae were generally and heavily stained. These r e s u l t s i n d i c a t e that the c e l l s of some t i n t i n n i d species are s u r p r i s i n g l y and r e l a t i v e l y impermeable to some dissolved substances. T i n -t i n n i d s i n t h i s area are extremely euryhaline, but the c o n t r i b u t i o n of the r e l a t i v e impermeability of the plasmalemma to t h i s e u r y h a l i n i t y i s not known. Unlike some estuarine and brackish-water c i l i a t e s , t i n t i n n i d s have no apparent vacuole to a i d i n osmoregulation. Centric diatoms dominate the larger phytoplankton i n the Vancouver area 59 except i n mid-summer. The only diatoms seen ins i d e t i n t i n n i d s from f i e l d samples were occasional small pennate diatoms, probably of the genus N i t z s c h i a ; and s i n g l e c e l l s of the small c e n t r i c diatom Skeletonema costatum. Even small diatoms usually occur i n chains of c e l l s and consequently are too large f o r t i n t i n n i d s to ingest. Diatoms thus form part of planktonic food chains which overlap i n s i z e those containing t i n t i n n i d s , but which are inde-pendent of them. However, l a r g e r zooplankton eat both diatoms and t i n t i n n i d s . The l a r g e s t food items contained i n Favella serrata and Eutintinnus l a t u s are always d i n o f l a g e l l a t e s of various species. Estimates of the ' e l e c t i v i t y ' or apparent d i f f e r e n t i a l s e l e c t i o n of food items of various types are given i n Sections 4a and 4b. The R o t i f e r species (mostly of the genus Synchaeta), copepod and barnacle n a u p l i i , and gastropod larvae were apparently s i m i l a r to t i n t i n n i d s i n being generally unselective predators (Table 2). The large h o l o t r i c h c i l i a t e Prorodon sp. was the only non-tintinnid c i l i a t e y which did not appear to eat laboratory phytoplankton. However, despite the f a c t that t h i s species i s quasisymbiotic (Blackbourn e t . a l . , 1973), on one occasion i t contained several 9.5 /im diameter polystyrene l a t e x p a r t i c l e s and one T i n t i n - nopsis nanaceell. 60 a) Accumulation experiments ( i i ) Quantitative Results General Tables3 to 24 show the r e s u l t s of a v a r i e t y of feeding experiments and associated experiments of a s i m i l a r nature. In most cases experiments were done with modified natural samples. Such samples usually contained a pre-ponderance of 1 or 2 species of t i n t i n n i d and a few i n d i v i d u a l s of several other species. Since there i s no other quantitative data on t i n t i n n i d feed-ing, the r e s u l t s of a l l species encountered i n an experiment have been given i n Tables 3 to 24 however small the number of c e l l s involved. The simplest experimental r e s u l t s are shown before the more complex; both i n terms of objectives e.g. comparisons of the r e l a t i v e s e l e c t i o n of several prey species, or estimating only uptake rates of simultaneous uptake and l o s s r ates; and also i n terms of the number of t i n t i n n i d species per experiment. In several s e l e c t i o n experiments, the t i n t i n n i d s i n some sections died or were moribund, leaving the r e s u l t s incomplete. The method of 'scoring' used was one i n which the number of c e l l s of a p a r t i c u l a r food type accumulated i n each t i n t i n n i d c e l l since the s t a r t of the experiment, or since the l a s t sub-sample of t i n t i n n i d s , was counted. Unless otherwise noted, the sub-samples were not preserved and counted l a t e r (see Methods Section), so there i s i n most cases an unavoidable l a g i n the timing of sub-samples taken from otherwise comparable treatments. For the most of the experiments, a one-way anaalysis of variance was performed together with Scheffe's t e s t f o r multiple comparisons with unequal sample sizes amongst a l l ' l e v e l s ' i n the experiment. The l e v e l s are identii-f i e d i n each Table by a small ringed number. To obtain homogeneity of 61 variance amongst the mostly small and highly v a r i a b l e l e v e l s , a log 10 trans-formation was made of each food c e l l count, a f t e r 1.0 had been added to each value to avoid zero values. In each Table, the r e s u l t of the one-way analysis of variance i s represented by the F-value and i t s s i g n i f i c a n c e l e v e l . Only those (Scheffe's) comparisons between l e v e l s which had a p r o b a b i l i t y of l e s s than 0.05 of being the same, are shown i n the Tables. L i t t l e emphasis has been placed on d i r e c t l y comparing or condensing r e s u l t s on the same species of t i n t i n n i d from d i f f e r e n t experiments, p a r t i c u l a r l y i f long periods of time separate the experiments. This i s because of the apparent s e n s i t i v i t y of t i n t i n n i d s to environmental factors other than food, and the large between-and within - experiment v a r i a b i l i t y i n feeding r a t e s . The probable causes of v a r i a b i l i t y i n t i n t i n n i d feeding rates are discussed l a t e r i n t h i s Section. Apparently r e c u r r i n g phenomena are discussed although the Scheffe's com-parisons i n any one experiment on which they are based may not show a s i g n i f i -cant d i f f e r e n c e . One predator type and one prey type Tables 3 to 6 show the r e s u l t s of simple experiments involving only one predator type and one prey type but with more than one concentration of food used. The feeding rates (in ml/hr/tin) of Eutintinnus tubulosus and T i n t i n - nopsis parvula were extremely low (<0.0001) i n these experiments even at high 3 food concentrations, i n one case i n excess of 73 x 10 c e l l s / m l . This was probably because the experiments la s t e d more than 3 hours, by which time a possible 'steady-state' had been reached between food eaten and food l o s t or digested. The c a l c u l a t i o n : accumulated food c e l l s / t i m e d i d not then accurately represent a feeding rate. Table 3 shows that there was no s i g n i f i c a n t d i f f e r e n c e between the c e l l s 62 T A B L E 3. Eutintinnus tubulosus feeding on Isocrysis galbana at two concentrations. Temperature 18°C; Salinity 11.5^. Duration (hrs) Numbers of prey-examined Numbers of prey/ predator Percentage Volume Numbers of prey of prey/ of digested predator pr e y / ml . F R nos . /hr / Tin . F R m l / h r / Tin . 3.7 5 15.4 + 5.1 21 770 53.400 4.2 0.00007 3.0 6 21.3 +9.0 27 "" 1065 73.600 7.1 0.00009 One-way A N O V A F value Significant Comparisons (Scheffes Test) with a significant difference at .0 5 level 1.28 No. (.05) N i l T A B L i E 4: Eutintinnus tubulosus feeding on Monochrysis lutheri at two concentrations. Temperature 17°C; Salinity 9.5&. Duration (hrs) Number s of prey examined Numbers of prey/ predator Percentage of prey digested Volume of prey/ predator Numbers of prey/ ml FR nos . /hr / T i n . FR m l / h r / T i n . 4.0 6 16.8 +6.9 15 840 4, 600 4.2 0.0009 5.0 7 17.7 + 5.9 14 885 21, 000 3.5 0.00016 One -way F value A N O V A Significant Comparisons (Scheffes test) with a significant difference at .05 level 0.02 No (.0 5) Nil 63 accumulated per t i n t i n n i d by Eutintinnus tubulosus i n 3 to 3.7 hours at two high but very d i f f e r e n t concentrations of Isochrysis galbana. Likewise, i n Table 4 E. tubulosus accumulated a s i m i l a r number of Monochrysis l u t h e r i c e l l s at two very d i f f e r e n t concentrations i n an experiment l a s t i n g 4 to 5 hrs. Obviously some sort of n u t r i t i o n a l steady-state had been reached i n both the experimental r e s u l t s shown i n Table 3 and 4, and i t i s also possible that 4,600 M. l u t h e r i c e l l s / m l i s approximately an optimal food concentration (OFC-see Methods Section) for E_. tubulosus. The very high food concentrations i n Table 3 were probably i n h i b i t o r y to the feeding of E_. tubulosus. Again, i n Table 5 i t can be seen that E_. tubuolsus accumulated i t s maximum prey c e l l number i n l e s s than 5.3 hours even at 9,000 c e l l s / m l of M. l u t h e r i , and that t h i s maximum number was quite s i m i l i a r to that i n Tables 3 and 4. I t i s i n -ter e s t i n g that the percentage of prey c e l l s undergoing d i g e s t i o n was f a i r l y s i m i l a r (14 to 31%) i n a l l 3 experiments. The number of c e l l s of Monochrysis l u t h e r i accumulated by Tintinnopsis  parvula can be seen i n Table 6 to be s i m i l a r i n 3 very d i f f e r e n t concentra-. tions of food i n experiments l a s t i n g 5 to 6 hours. The number accumulated at 23 to 28 hours a f t e r the s t a r t was also s i m i l a r i n the three food concen-t r a t i o n s , and was much lower than at the 5 to 6 hours' check. This i s probably due to reduced feeding a c t i v i t y caused by some p h y s i o l o g i c a l s t r e s s , as can be seen from the f a c t that T_. parvula was l e s s a c t i v e i n the l a t e r check. The d i g e s t i o n rate was apparently l e s s a f f e c t e d by t h i s p o s s i b l e s t r e s s . A much higher proportion of M. l u t h e r i c e l l s were undergoing diges-t i o n by T_. parvula i n the l a t e r check than i n the e a r l i e r check. From Tables 5 and 6 i t seems that J_. parvula has an accumulated food maximum of about 20 c e l l s of M. l u t h e r i (at 9 or 10°C.) and that i t s OFC i s below 7000 c e l l s / m l of M. l u t h e r i . 64 T A B L E 5. Eutintinnus tubulosus feeding on Monochrysis lutheri at three concentrations. Temperature 16°C; Salinity 28%,,. Duration (hrs.) Numbers of prey examined Numbers of prey/ predator Percentage of prey dige sted Number s of prey/ ml. FR nos . /hr / T i n . FR m l / h r / Tin. 6.3 n 25.0 + 6.6 20- 9. 000 4.0 0.00044 5.6 13 29.0 + 10.1 28 18.000 5.2 0.00028 5.3 7 25.7 + 10.2 31 72, 000 4.9 0.00006 One-way A N O V A Comparisons (Scheffes Test) with a significant difference F-value Significant at .05 level.  0.64 No (0.0 5) N i l T A B L E 6. Tintinnopsis parvula feeding on Monochrysis lutheri at three concentrations. Two samples taken from each. Temperature 9 °C; Salinity 24J&. Duration (hrs) Numbers of prey examined Number s of prey/ pr edator Percentage of prey dige sted Numbers of p r e y / m l . FR nos . /hr / Tin . FR m l / h r / T i n . T i n . condition T i n . Dividing 6.5fT) 9 21.3+ 6.5 6 14, 000 3.3 0.0002 Active N i l 28.0^. 3 6.7 + 4.7 50 14, 000 Slow N i l ~ 6 . 0 © 16 18.6+ 7.7 6 29. 000 3.1 0.0001 Active N i l 2 8 .00 10 5.4 + 3.6 35 29. 000 Slow N i l . 5-°© 10 28.7 + 12.1 18 73, 500 5.7 0.00007 Active no granules 2 2 3 . 0 © 11 6.6 + 6.2 64 73, 500 ™* *" No or few granules N i l One-way A N O V A F-value Significant Comparisons (Scheffes Test) with a significant difference at .05 level 10.10 Yes (.01) 1/4, 1/6. 3/4. 3/6. 4/5 65 From Tables 3 to 6 i t can be seen that optimal food concentrations ( i . e . optimal f o r maximum feeding rate) f o r the medium-sized t i n t i n n i d s Eutintinnus tubulosus^and Tintinnopsis parvula seem to be l e s s than 4,000 c e l l s / m l of 3 small prey species of about 50 um volume. Steady-state feeding rates under these conditions were reached i n l e s s than 3 hours and the equivalent maximum 3 number of p r e y / t i n t i n n i d was about 20 to 30 or about 1,000 to 1,500 um . Table 7 shows the r e s u l t of the only f a c t o r i a l design among the accum-u l a t i o n experiments. I t involved Tintinnopsis subacuta feeding on D u n a l i e l l a  t e r t i o l e c t a . Samples were taken at three times a f t e r the s t a r t : at 0.66 hrs, 1.5 hrs and 6.0 hrs; at three temperatures: 10.0, 13.2 and 19.8 C; and at three concentrations of food: 1,750, 8,700 and at 43,400 c e l l s / m l . The sampling was staggered i n the same order and for the same length of time across temperatures and food l e v e l s at each time check; and samples were immediately added to formaldehyde to a f i n a l concentration of about 2%. Ad-herence to t h i s procedure made possible the removal of as much sampling bias as p o s s i b l e . The variance across the 3 food l e v e l s was by far the most s i g -n i f i c a n t , followed by that i n the i n t e r a c t i o n between temperature and food l e v e l . Variance across the three l e v e l s of time and i n the i n t e r a c t i o n s be-tween a l l 3 factors was s i g n i f i c a n t at the .05 l e v e l . Variance across the l e v e l s of temperature and between time and temperature was not s i g n i f i c a n t at the .05 l e v e l . I t might be true to say that food l e v e l i s the most important v a r i a b l e of these to T_. subacuta, followed by temperature and time. T_. subacuta would never be subjected to a temperature as high as 19.8 C i n a natural s i t u a t i o n ; nor i s i t l i k e l y to be presented with as many as 43,400 c e l l s / m l of one species of natural food. I t i s i n t e r e s t i n g that the percen- \ tage of semi-digested c e l l s was greater a t 19.8 C than at the lower temper-atures, e s p e c i a l l y at the lower food l e v e l s . I t w i l l also be seen i n other 66 T A B L E 7. Tintinnopsis subacuta feeding on Dunaliella tertiolecta (ZIP um ) at three temperatures and three food levels. Formalin used. Salinity 9J&, dark. Number Number Percentage Volume Number ' FR FR Duration predator s Temp. prey/ prey prey/ of nos./hr/ ml/hr/ (hrs) Predator examined °C predator digested predator prey/ml Tin. Tin. 0.66 T. subacuta 8 10.0 3.4+ 2.6 Nil 714 1. 750 5.2 0.0030 1.5 T. subacuta 2 10.0 1.5+0.7 Nil 315 1. 750 1.0 0.00057 6.0 T . subacuta 3 10.0 4.3+3.1 62 903 1. 750 0.66 T. subacuta 12 13.2 7.5+3.5 Nil 1. 575 1. 750 11.4 0.0065 1.5 T. subacuta 3 13.2 7.3+ 4.5 Nil 1. 533 1. 750 4.9 0.0028 6.0 T . subacuta 3 13.2 3.3+1.6' 50 693 1. 750 0.66 T . subacuta 9 19.8 1.0+ 1.4 Nil 210 1. 750 1.5 0.00087 1.5 TV subacuta 4 19.8 3.5+2.9 79 735 1. 750 2.3 0.0013 6.0 T. subacuta 6 19.8 2.5+6.1 80 525 1. 750 0.66 T. subacuta 8 10.0 6.6+4.3 11 1, 386 8, 700 10.0 0.0012 1.5 T. subacuta 5 10.0 12.8+ 4.8 . 13 2, 688 8. 700 8.5 0.0010 6.0 T . subacuta 7 10.0 12.0+8.3 18 2, 520 8, 700 - - -0.66 T. subacuta 4 13.2 7.0+4.2 11 1. 470 8, 700 10.6 0.0012 1.5 T. subacuta 4 • 13.2 13.0+7.9 Nil • 2, 730 8. 700 8.7 0.0010 6.0 T. subacuta 3 13.2 8.0+ 5.6 83 1. 680 8. 700 0.66 T. subacuta 6 19.8 9.3+8.6 14 1. 953 8. 700 14.1 0.0016 1.5 T. subacuta 3 19.8 24.0+ 6.9 54 5. 040 8, 700 16.0 0.0018 6.0 T. subacuta 9 19.8 15.9+9.0 68 3. 339 8, 700 0.66 T. subacuta 5 10.0 23.4+ 7.2 10 4, 914 43, 400 35.5 0.00082 1.5 T. subacuta 9 10.0 24.3+ 6.6 11 5, 103 43. 400 16.2 - 0.00037 6.0 T . subacuta 3 10.0 16.0+ 5.3 50 3. 360 43, 400 0.66 T. subacuta 9 13.2 17.6+6.6 4 3. 696 43, 400 26.7 0.00061 1.5 T. subacuta 7 13.2 19.9+8.3 4 4. 179 43, 400 13.3 0.00031 6.0 T. subacuta 4 13.2 32.5+14.8 45 ' 6, 825 43, 400 . . . 0.66 T. subacuta 8 19.8 11.1+ 8.0 11 2. 331 43. 400 16.8 0.00039 1.5 T. subacuta 3 19.8 19.7+3.1 42 4. 137 43, 400 13.1 0.00030 6.0 T. subacuta 3 19.8 26.3 + 16.2 61 5. 523 43. 400 Analysis of Variance for 3x3x3 factorial on log-transformed values +1.0  Interaction Time Temperature Time x Temperature Food level Time x Food level Temperature x Food level Time x Temperature x Food level 2.42 F-value Significant 4.58 yes (.05) 0.45 ho 1.41 no 70.16 yes (.01) 2.71 yes (.05) 3.78 yes (.01) 2.42 yes (.05) 67 Tables that digestion rate appears to be more d i r e c t l y affected by temper-ature than does feeding rate. Two or more predator types and one prey type In Table 8 Tintinnopsis parvula and Tintinnopsis c y l i n d r i c a were exposed to moderate numbers of Monochrysis l u t h e r i at only one l e v e l (7,000 cells/ml) i n dim l i g h t and also i n darkness, both at 10 C. This experiment was much shorter than those shown inTable 3 to 6 and t h i s i s r e f l e c t e d i n the small percentage of M. l u t h e r i c e l l s undergoing di g e s t i o n ( 10% u n t i l a f t e r 1 hour). There was no s i g n i f i c a n t d i f f e r e n c e between the accumulated c e l l number of T_. parvula and that of T_. c y l i n d r i c a i n darkness or i n dim l i g h t (also see Methods Section). The maximum i n i t i a l feeding rate of the two species i n Table 8 - about 70 c e l l s / h r / t i n t i n n i d or equivalent to about 1% ml/hr - was fas t e r than those shown i n Tables 3 to 6 and was the f a s t e s t feeding rate seen i n these accumulation experiments. However, the maximum accumulation of prey c e l l s was s t i l l only about 20 per t i n t i n n i d (see above). In Table 9 two t i n t i n n i d species, Eutintinnus tubulosus and Helic o s t o -mella k i l i e n s i s , are compared when feeding on Monochrysis l u t h e r i at 3 very dense concentrations. The accumulated food c e l l numbers are again t y p i c a l of those i n Table 3 to 6 i n 41,400 ce l l s / m l and i n 107,000 c e l l s ^ m l , and only lower and s i g n i f i c a n t l y d i f f e r e n t from these at 213,000 c e l l s / m l . The contact rates (see Section 4b) of these species were concurrently observed i n t h i s experiment (Table 9), and t h i s rate declined i n H. k i l i e n s i s through a 5-fold increase i n prey c e l l concentrations. The contact rate of E. tubu-losus^ increased r e l a t i v e l y l e s s than did the prey c e l l concentration i n the samples (Table 9). These very high l e v e l s of food c e l l s (or t h e i r associated metabolites) were obviously close to the l e v e l at which the normal feeding 68 T A B L E 8. T i n t i n n o p s i s p a r v u l a a n d T. c y l i n d r i c a f e e d i n g o n M o n o c h r y s i s l u t h e r i . i n d i m l i g h t a n d i n d a r k n e s s . T e m p e r a t u r e 1 0 °C; S a l i n i t y 2.3%,. N u m b e r N u m b e r P e r c e n t a g e V o l u m e N u m b e r F R F R D u r a t i o n p r e d a t o r s p r e y / p r e y p r e y / o f n o s . / h r / m l / h r / I l l u m i -( h r s ) P r e d a t o r e x a m i n e d p r e d a t o r d i g e s t e d p r e d a t o r p r e y / m l T i n . T i n . n a t i o n 0 .25 T . p a r v u l a 6 15 .2+6.6 N i l 760 7. 000 60.8 0 .0087 D a r k 0.25 T . c y l i n d r i c a 5 18.4+ 9.2 N i l 920 7, 000 7 3 . 6 0.011 D a r k 0 .75 T , p a r v u l a 8 13 .9+3 .2 9 690 7 ,000 18 .5 0 .0026 D a r k 0 .75 T . c y l i n d r i c a 1 11.0 N i l 550 7. 000 14.6 0 .0021 D a r k 1.25 T . p a r v u l a 8 16.4+ 6.0 5 820 7. 000 13.1 0 .0019 D a r k 1.25 T . c y l i n d r i c a 1 10.0 N i l 500 7. 000 8.0 0 .0011 D a r k 0.50 T . p a r v u l a 5 13.0+ 11.0 N i l 650 7, 000 26.0 0 .0037 D i m 1.0 T . p a r v u l a 6 20.8+ 5.5 10 1040 7, 000 20.8 0 .0030 D i m 1.0 " T . c y l i n d r i c a 1 13.0 45 650 7 , 0 0 0 13.0 0 .0019 D i m 1.5 T . p a r v u l a 1 19.0 58 950 7. 000 13.0 0 . 0 0 1 9 D i m 1.5 T . c y l i n d r i c a 2 16.5+ 7.5 28 825 . 7 , 0 0 0 11.0 0 .0016 D i m O n e - w a y A N O V A C o m p a r i s o n s ( S c h e f f e s T e s t ) w i t h a s i g n i f i c a n t d i f f e r e n c e F - V a l u e S i g n i f i c a n t a t . 0 5 l e v e l  0.98 N o N U T A B L E 9. Eutintinnus tubulosus and Helicostomella kiliensis feeding on Monochrysis lutheri at three concentrations. Temperature 16°C; Salinity 15^,. Duration Number predators Number Percentage Volume Number FR FR prey/ prey prey/ of nos./hr/ ml/hr/ Tins (hrs ) Predator examined predator dige stei d predator prey/ml Tin. Tin. dividing 8 . 5 © E. tubulosus 3 15.0+ 8.0 30 750 41,400 1.8 0.00004 Nil 8 . 5 © H. kiliensis 3 14.7+ 2.6 34 735 41, 400 1.7 0.00004 1 8 . 0 0 E . tubulosus 6 22.0+ 3.0 17 1100 107, 000 2.8 0.00002 Nil 8.0 H. kiliensis 1 3.0 . 100 150 107, 000 Nil E . tubulosus 1 9.0 67 450 213, 000 1.3 Nil 7.0 H. kiliensis 8 5.5+ 9.3 59 275 213, 000 0.8 4 One-way ANOVA F-value Significant Comparisons (Scheffes Test) with a significant difference at .0 5 level 7.36 yes (.01) 2/4, 3/4 CONTACT RATES Predator Number predators examined Total Duration (sees) Total Contacts Ingested Number prey/mi Contacts/ ' minute CR ml/hr/ Tin. FR ml/hr/ Tin. E . tubulosus 1 50 • 9 ' Nil 4L 400 10 . 0.015 Nil H. kiliensis 1 20 8 Nil 41, 400 24 0.035 Nil E . tubulosus 4 159 39 Nil 107, 000 14.7 0.0082 Nil H. kiliensis 3 236 62 N i l - 213, 000 15.8 0.0045 Nil 70 behaviour of these t i n t i n n i d species breaks down. These l e v e l s are probably 5 to 10 times higher thanbhigh f i e l d concentrations of food c e l l s of t h i s s i z e . The low l e v e l of feeding a c t i v i t y i n H. k i l i e n s i s i n 213,000 prey c e l l s / m l was, however, associated with a high l e v e l of reproductive a c t i v i t y , and 4 of the 8 sampled i n d i v i d u a l s showed early signs of c e l l d i v i s i o n (oral Ahlage). In Table 10 the r e s u l t s are shown of an experiment i n which 5 species of t i n t i n n i d were exposed to four concentrations of D u n a l i e l l a t e r t i o l e c t a . The highest concentration (19,400 cells/ml) was from a month-old stock c u l -ture and the lower concentrations were from stocks le s s than 1 week ol d . The age of the culture made no d i f f e r e n c e i n t h i s experiment. The percentage of prey being digested had a s i m i l a r range i n a l l species and was 18 to 39% i n Tintinnopsis subacuta, 17 to 36% i n T_. parvula and 9 to 40% i n Stenosomella  ventricosa . , No prey c e l l s were digested i n S_. n i v a l i s and 100% were digested by the rare t i n t i n n i d Ptychocyclis acuta. T_. subacuta and S_. ventricosa had s i m i l a r feeding rates (about 0.0020 ml/hr), with those of the l a t t e r being a l i t t l e higher, but not s i g n i f i c a n t l y so. The optimal food concentration (OFC) was between 2,000 and 3,900 c e l l / m l f o r T_. subacuta and S^. ventricosa or 0.37 to 0.72 ppm by volume. The feeding r a t e of Tintinnopsis parvula was only 1/4 or 1/5 of that of the l a t t e r two species at the same food c e l l concentration; and the smallest of the species Stenosomella n i v a l i s , had an accumulation of prey c e l l s only 1/10 of that of T_. subacuta i n t h i s experiment. The r e s u l t s shown i n Tables 8 to 10 of the accumulation and feeding rates of more than one t i n t i n n i d species, are an extension of those of Tables 3 to 6 i n i n d i c a t i n g that the average number of c e l l s accumulated/tintinnid was about 20oM. 3lu6Reri?; and that optimal food concentrations of M. l u t h e r i were about 71 T A B L E 10. Various tintinnid species feeding on "new" and "old" cultures of Dunaliella  tertiolecta at four concentrations. Temperature 14°c; Salinity 28^. Duration (hrs) Predator Number Number Volume of predators prey/ pred/prey examined predator (um3) Percentage FR FR prey Number nos/hr/ ml/hr/ digesting prey/ml Tin. Tin. Age of prey stock culture 0 + 1.8Q Tintinnopsis subacuta 0 + 2 .0^ Tintinnopsis subacuta 0 + 2.25 Tintinnopsis ® subacuta 0 + 2. 5jv Tintinnopsis 0 + 2.&£> Tintinnopsis subacuta 0 + 2.00 Tintinnopsis parvula 0 + 2.25 Tintinnopsis >Z> parvula 0 + 1.8 Stenosomella ventricosa 0 + 2.0 Stenosomella © ventricosa 0 + 2.25 Stenosomella © ventricosa 0+2 .5 Stenosomella ventricosa . 0 + 2.8. Stenosomella <s ventricosa 0 + 2.25, Stenosomella nivalis 0 + 2.5 Stenosomella nivalis 0 + 2.8 Stenosomella nivalis 0 + 2.25 Ptychocyclis acuta 10 3.6+3.4 702 14.0+ 9.8 2730 19.9+12.3 3881 T*) c ~ ~ w - subacuta 12 21.9+12.3 4271 11 25.1+ 8.9 4718 3.0+3.3 18.0 N i l 2.0 N i l 2.0 585 5.5+ 6.0 1073 3510 15.7+ 5.9 3062 30.3 + 9.3 5909 15.0+ 9.9 2925 46.7+14.2 8780 N i l 390 N i l 390 39 31 18 28 17 17 36 11 26 40 12 9 N i l N i l 100 2,000 3. 900 7, 800 15. 600 19.400 3.900 7. 800 2,000 3. 900 7. 800 15. 600 19. 400 7. 800 15. 600 19.400 7, 800 2.1 7.0 8.8 8.7 8.9 13 2.4 10.0 7.9 13.5 6.0 16.5 0.0011 0.0018 0.0011 0.0006 0.0005 0.0004 0.0003 0.0051 0.0020 0.0017 0.0004 0.0009 0.8 0.00005 0.9 0.00011 4 days 4 days 4 days 4 days 1 month 4 days 4 days 4 days 4 days 4 days 4 days 1 month 4 days 4 days 1 month 4 days One -way ANOVA F-value Significant 7.76 yes (.01) Comparisons (Scheffes Test) with a significant difference at .05 level  1/3. 1/4. 1/5. 1/9, 1/11. 6/11 72 2 or 3,000 per ml. I t i s i n t e r e s t i n g that the estimated feeding rates of t i n t i n n i d s sampled at 1 hour i n Table 8 were considerably lower than the i n i t i a l rates at 0.25 hours, even though the number of semi-digested c e l l s at 1 hour was s t i l l very low (<10%). Therefore, i t seems that here ingestion slowed well before the di g e s t i v e capacity of the c e l l s was saturated, but the accumulated prey numbers/tintinnid at 0.25 hours were close to the maxi-mum. I t seems that the t i n t i n n i d s were apparently near saturation i n terms of numbers at 0.25 hours (see end of t h i s Section). Table 10 shows that the smaller t i n t i n n i d species accumulated many fewer D u n a l i e l l a t e r t i o l e c t a than d i d the larger species. The OFC of T_. subacuta on t h i s prey species was about 4,000/ml. D i f f e r e n t i a l predation and s e l e c t i o n experiments One predator type and two or more prey types In the following Tables Ivlev's e l e c t i v i t y index has been used as a measure of comparative ' c a t c h a b i l i t y ' or of d i f f e r e n t i a l predation and not as an index of prey s e l e c t i o n (and see Section 4c). Tables 11 to 13 show the r e s u l t s of one predator exposed to 2 or more prey types simultaneously. In the experiment shown i n Table 11 Tintinnopsis parvula was exposed f o r about 1 hour to Isoselmis sp. s i n g l y at 9,500 c e l l s / m l , and to Isoselmis  sp. plus Monochrysis l u t h e r i at 9,500 and at 14,000 c e l l s / m l , r e s p e c t i v e l y . The number of Isoselmis sp. accumulated/tintinnid was no d i f f e r e n t i n the two s i t u a t i o n s , and was much lower than the number of M. l u t h e r i accumulated. The e l e c t i v i t y index indicated that a disproportionately large number of M. l u t h e r i , and a disproportionately small number of Isoselmis sp. were accum-ulated from the prey food mixture, but t h i s shows only d i f f e r e n t i a l predation. Although Isoselmis sp. i s a l i t t l e l a r g e r , and usually moves f a s t e r than 73 T A B L E 11. Tintinnopsis parvula feeding on Isoselmis sp. and Monochrysis lutheri. Temperature 1 6 ° C ; Salinity 15&. Duration (hrs) Number predators examined Prey Number prey/ predator Volume prey/ predator Number D r e y / m l F R N o s / h r / T i n F R m l / h r / T in E l e c -tivity Index O . 8 0 7 Isoselmis sp. 1.9+ 1.9 140 9. 500 2.4 0.00025 © © 4 Isoselmis sp. and M . lutheri 1.5+ 2.1 . 15.0+7.8 113 750 -9. 500 14.000 1.4 13.6 0.00014 0.0010 -.63 +.28 Total 16.5 863 23, 500 15.0 0.00063 One-way A N O V A F-value Significant 11.56 yes (.01) Comparisons (Scheffe's Test) with a significant difference at .0 5 level  1/3. 2/3 T A B L E 12. Tintinnopsis subacuta feeding on Eutreptiella sp. (500 um 3 ) and Isochrysis galbana (45 u m 3 ) . Temperature 1 4 ° C ; Salinity 9&. Duration (hrs) Number predator s examined Prey Number prey/ predator Vol . prey/ predator (um3) Number p r e y / m l F R N o s / h r / T in . F R m l / h r / T in . E l e c -tivity Index 1 1 . 3 @ 8 Eutreptiella only 16.4+ 8.3 8, 200 "4.4 x 10 3 1.5 ? minimal 0.0003 ? minimal 12.5 © 11 I. galbana only 26.3+12.2 1, 200 66 x 10 3 2.1 minimal 0.00003 ? minimal . . . 13 Eutreptiella sp. and 18.3+ 7.6 9. 100 4.4 x 10 3 1.6 ? minimal 0.0004 minimal + .86 © 1. galbana 4.5+ 8.2 203 66 x 10 3 0.4 minimal 6 x 10"6 minimal - .65 Total 22.8 9. 30 3 70.4x 10 3 2.0 One-way A N O V A F-value Significant Comparisons (Scheffe's Test) with a significant difference at .0 5 level 25.11 yes (.01) 1/4. 2/4. 3/4 74 M. l u t h e r i i t s l i a b i l i t y to predation was lower i n th i s experiment. Tables 12 and 13 show the r e s u l t s of Tintinnopsis subacuta alone exposed to E u t r e p t i e l l a sp. and another prey species i n r e l a t i v e l y long experiments. In Table 12 the r e s u l t s are seen of an experiment l a s t i n g 11.3 to 12.5 hours with T_. subacuta exposed to E u t r e p t i e l l a sp. s i n g l y at 4,400 c e l l s / m l ; to Isochrysis ga-lbana at 66,000 c e l l s / m l ; and to a dense mixture of the two species at 4,400 c e l l s and 66,000 c e l l s / m l r e s p e c t i v e l y . There was no s i g -n i f i c a n t d i f f e r e n c e between the number of accumulated c e l l s (steady-state balance) per t i n t i n n i d i n E u t r e p t i e l l a sp. only; i n I_. galbana only; or of E u t r e p t i e l l a sp. i n the mixed s i t u a t i o n . As i n some previous experiments the maximum of prey c e l l s / t i n t i n n i d was on average between 16 and 25 despite a 10-fold d i f f e r e n c e i n prey volume. I t i s also noteworthy that proportionately more E u t r e p t i e l l a sp. than I_. galbana were accumulated when both were pre-sented s i n g l y . However, the number of I_. galbana accumulated over 11.5 hours i n the mixed s i t u a t i o n was s i g n i f i c a n t l y reduced to about 1/6 of that of the number accumulated when exposed to I_. galbana s i n g l y . An i n d i v i d u a l T_. suba- cuta with a new l o r i c a i n E u t r e p t i e l l a sp. contained merely 6 prey items a f t e r up to 11.3 hours, whereas i n the mixed prey s i t u a t i o n a s i m i l a r i n d i v i -dual with a new l o r i c a contained 40 E u t r e p t i e l l a sp. and 30 I_. galbana i n up to 11.5 hours. Therefore, i t seems that the great v a r i a b i l i t y i n i n d i v i d u a l t i n t i n n i d d feeding performance seen i n a l l these experiments can be found even amongst i n d i v i d u a l s with a r e l a t i v e l y s i m i l a r recent p h y s i o l o g i c a l h i s -tory. The estimated feeding rates i n Table 12 are extremely low due to the fa c t that a f t e r 11 to 12 hours, the t i n t i n n i d s were probably i n some sort of steady-state with regard to the uptake of these food items. Table 13 shows the r e s u l t s of another experiment of t h i s type, but also T A B L E 13. Tintinnopsis subacuta feeding on Eutreptiella sp. (500 um 3 ) . Isochrysis galbana (45 um 3 ) . and Dunaliella tertiolecta (210 unv*). . ..Temperature 1 4 ° C ; Salinity 9&. Number Duration predators (hrs) examined P r e y Number Volume prey/ .. prey/ . Number predator predator prey/ml FR FR E l e c -Nos /hr / m l / h r / tivity Tin Tin Index 6.0@ ® 6.3 © 6.60 Eutreptiella and D. tertiolecta Total D. tertiolecta only D. tertiolecta and I_. galbana 16.0+2.8 8.0 x 103 6.6+ 2.0 1. 390 22.6 9.7+1.8 2.7+2.1 46.7+21.9 9, 390 2, 040 570 2, 100 1, 500 4. 400 5. 900 2.7 min. 0.0018 min. +.47 1.1 min. 0.00025 -.44 4,400 1.5 min. 0.00034 min. - - -4.400 0.41 min. 0.00009 min. -.07 66,000 7.1 min. 0.00011 min. +.004 Total 49.4 2, 670 70,400 One-way A N O V A F-value Significant 24.24 yes (.01) Comparisons (Scheffe's Test) with a significant difference at .05 level. 1/4, 2/5. 3/4, 3/5, 4/5 76 inc l u d i n g the f l a g e l l a t e D u n a l i e l l a t e r t i o l e c t a . Tintinnopsis subacuta was exposed to p_. t e r t i o l e c t a s i n g l y at 4,400 c e l l s / m l ; to E u t r e p t i e l l a sp. (1,500/ml) plus D. t e r t i o l e c t a (4,400/ml); or to D. t e r t i o l e c t a (4,400./ml) plus Isochrysis galbana (66,000/ml). A l l t i n t i n n i d s i n the other possible single-food'situations i n t h i s experiment; e.g. with E u t r e p t i e l l a sp. and I_. galbana, died from unknown causes during the experiment. More c e l l s of p_. t e r t i o l e c t a were accumulated per t i n t i n n i d when presented s i n g l y than when presented together with I_. galbana or E u t r e p t i e l l a sp. , although the di f f e r e n c e was not s i g n i f i c a n t i n the l a t t e r case. Despite these r e a l numer-i c a l d i f f e r e n c e s , p_. t e r t i o l e c t a was apparently accumulated only s l i g h t l y l e s s than i n the proportion i n which i t occurred, i n the very dense mixture with J . . galbana ( e l e c t i v i t y of -.07); but was accumulated much less than proporr". t i o n a t e l y i n the mixture with E u t r e p t i e l l a sp. ( e l e c t i v i t y of -,44)jbut as i t can be seen from Table 13, t h i s apparent d i f f e r e n c e i s due to the peculiar nature of Ivlev's formula. The s l i g h t l y decreased accumulation of p_. t e r t i o - l e c t a i n either of the mixed-prey s i t u a t i o n s compared with the single-prey s i t u a t i o n i s a s i m i l a r r e s u l t to that of I.galbana when exposed with E u t r e p t i e l l a sp. inTTable 12 and has no obvious behavioural explanation. It seems as though T_. subacuta cannot (or does not) accumulate more of the most e a s i l y caught item (e.g. E u t r e p t i e l l a sp. - i n Table 12), when i n a mixture than i t does i n single-prey s i t u a t i o n s , but that i t does take l e s s of the other prey species i n the mixture than s i n g l y . In Table 13 i t can be seen that-although about 47 I_. galbana and only about 16 E u t r e p t i e l l a sp. 3 were accumulated, prey volume/predator was about 8,000 pn for E u t r e p t i e l l a 3 sp. and only about 2,100 jam for I_. galbana. As both species are r a p i d l y digested by T_. subacuta, the t i n t i n n i d gained much more i n terms of biomass by feeding on E u t r e p t i e l l a sp. than on I_. galbana. T A B L E 14. Tintinnopsis subacuta and Tintinnidium mucicola feeding on Eutreptiella sp. (500 |im )and Isoselmis sp. (75 um3). Temperature 13°C; Salinity 12.5%. ,) Duration ( h r s ) P r e d a t o r Number predators examined P r e y Number Vol. prey/ FR FRX Elec-prey/ predator Number Nos/hr/ ml/hr/ tivity . predator (urn3) prey/ml Tin. Tin. Index; 0.6 T. subacutaQ 6 0.6 T. subacuta^ 6 © Eutreptiella and Isoselmis Total 5.3 + 2.0 2.0 + 2.0 2, 650 150 2. 800 3, 650 30,000 8.4 3.2 11.6 0.0023 0.00010 0.00034 +.71 -.53 0,6 T. mucicola^ 6 0.6 T. mucicola- 6 Eutreptiella and Isoselmis Total Nil 7.2 + 4.4 540 540 3, 650 30,000 Nil 11.4 Nil 0.00037 11.4 0.00037' -1.0 +.06 One -way ANOVA F-value Significant 3.45 No Comparisons (Scheffe's Test) with a significant difference at .0 5 level. • Nil 78 Two or more predators and two or more prey types In Table 14 the r e s u l t s are shown of a f a i r l y short experiment i n v o l -ving Tintinnopsis subacuta and the unusual species Tintinnidium mucicola. They were presented with only a mixture of E u t r e p t i e l l a sp. and Isoselmis sp. (cryptophycae) at 3,650 and 30,000 c e l l s / m l r e s p e c t i v e l y . After 0.6 hours T_. subacuta had accumulated disproportionately more of E u t r e p t i e l l a sp. than (3£ the much smaller and much more numerous Isoselmis sp. . On E u t r e p t i e l l a sp. T_. subacuta showed a f a s t estimated i n i t i a l feeding rate of 0.0023 ml/hr/ t i n t i n n i d . T_. mucicola d i d not eat E u t r e p t i e l l a sp. and accumulated more Isoselmis sp. than did T_. subacuta from the mixutre of prey types. However, due to the short feeding periods and small numbers of accumulated c e l l s , the l a t t e r were not quite s i g n i f i c a n t l y d i f f e r e n t at the .05 l e v e l . In Table 14 the e l e c t i v i t y indices are i n d i c a t i v e of r e a l trends but only by comparison with the other predator, and not by comparison with a single-prey s i t u a t i o n for the same predator. In the f i e l d sample from which the t i n t i n n i d s were taken f o r the l a t t e r experiment, the contact rate of T_. mucicola on a l l p a r t i c u l a t e material was a l i t t l e more than half that of T_. subacuta. It i s of s i m i l a r s i z e to the l a t t e r but moves more slowly (see Section 4b). Therefore', the f a c t that T_. mucicola may have accumulated more Isoselmis sp. than T_. subacuta cannot be ascribed to a fas t e r contact r a t e . Three of the s i x T_. subacuta shown i n Table 14 showed the ea r l y signs of c e l l d i v i s i o n . None of four T^ . subacuta taken from the same f i e l d sample as those used i n the experiment showed signs of c e l l d i v i s i o n , even though two of them had eaten one Tintinnopsis nana each. No T_. mucicola c e l l showed signs of c e l l d i v i s i o n i n the f i e l d sample or i n the experiment. It i s , therefore, perhaps possible that some T_. subacuta i n d i v i d u a l s i n the experiment had responded to the act of ingestion or the 79 presence of E u t r e p t i e l l a sp. i n t h e i r cytoplasm (none appeared to be digested) or i n the medium, and i n the space of 38 minutes had begun c e l l d i v i s i o n . The r e s u l t s i n Table 14 confirmed the apparent s e l e c t i v i t y of these two t i n t i n n i d species i n a f i e l d sample taken 4 days previously. In t h i s sample T_. subacuta contained mainly many c e l l s of a species of euglenoid (probably E u t r e p t i e l l a sp_.), and T_. mucicola contained only small reddish c e l l s 5-7 u^m i n diameter which were probably a species of cryptomonad. Similar observations were made on several other occasions. Table 15 shows the food c e l l s accumulated by Tintinnopsis subacuta, T_. nana, T_. rapa and Tintinnidium mucicola presented with Monochrysis l u t h e r i (13,000 cells/ml) and D u n a l i e l l a t e r t i o l e c t a (6,200 cells/ml) for 1 hour. Only T_- subacuta ate both prey species; M. l u t h e r i apparently rather more then proportionately compared to i t s abundance intthe medium, and p_. t e r t i o l e c t a rather l e s s so. However no sihgle=prey s i t u a t i o n was a v a i l a b l e for comparison. The average feeding rate of T_. subacuta on M. l u t h e r i was 26.5 c e l l s / h r / t i n t i n n i d or 0.002 m l / h r / t i n t i n n i d , a f i g u r e s i m i l a r to that on E u t r e p t i e l l a sp. i n Table 14. T_. rapa and T_j_ nana accumulated l e s s M^ l u t h e r i than did T. subacuta, and t h e i r contact rates and c e l l volumes (Table 1) are also much smaller than the l a t t e r ' s . T_. rapa and T_. nana ate no D_. t e r t i o l e c t a and they may be too small to ingest i t (Table 2). Tintinnidium  mucicola ate neither prey species on t h i s occasion for unknown reasons, a l -though most i n d i v i d u a l s did contain some old food items. Scheffe's test was not s i g n i f i c a n t for most of the comparisons between l e v e l s i n t h i s experiment due to the small sample s i z e s . Table 16 shows the r e s u l t s of an experiment i n which two t i n t i n n i d species of s i m i l a r s i z e and general 'searching r a t e ' Tintinnopsis subacuta T A B L E 15. Various tintinnid species feeding on Monochrysis lutheri (50 um ) and Dunaliella tertiolecta (200 um ). Temperature 16°C; Salinity 22^,. Number Number Volume FR F R E l e c -Duration predators prey/ prey/ Number Nos /hr / m l / h r / tivity' (hrs) Predator examined Prey predator predator prey/ml Tin . T i n . Index 0 + 1.0 Tintinnopsis M . lutheri 26.5+2.2 1325 13,000 26.5 0.0020 +.16 subacuta /rs 2 D . tertiolecta 2.0+ 2.0 400 6,200 2.0 0.0003 -.64 © 0+1.0 Tintinnopsis M . lutheri 6.5+0.7 325 13,000 6.5 0.0005 m » r apa © 2 D. tertiolecta Nil N i l 6, 200 N i l Nil --0+1.0 Tintinnopsis M . lutheri 1.8+2.4 . • 90 . 13,000 1.8 0.00013 — _ nana 0 4 • D. tertiolecta Nil N i l 6,200 N i l Ni l --0 + 1.0 Tintinnidium M . lutheri Nil Ni l 13,000 Nil Ni l mucicola 4 D. tertiolecta N i l N i l , 6,200 N i l Ni l One -way A N O V A F-value 1 Significant 5.76 Yes (.0 5) Comparisons (Scheffe's Test) with a significant difference at .05 level. 1/4 T A B L E 16. Tintinnopsis subacuta and Stenosomella ventricosa feeding on Eutreptiella sp. (450|im3), M o n o c h r y s i s l u t h e r i (50 ptm 3), and I sose lmis sp. (75|ima), singly and in combination. Duration (Hrs) Predator Number predator a examined Prey Number prey/ predator Percentage prey digested Volume prey/ predator Number . prey/ml F R nos /hr / Tin. F R m l / h r / T i n . Electivity Index (nos) 0 + 2.7 Tintinnopsis subacuta L 9 Eutreptiella sp. (only) 16.0+3.4 58 7200 1400 5.9 0.0042 --0 + 1.8 a 7 M . lutheri and 26.0+5.4 - • 1300 8000 14.4 0.0018 +.10 3 Isoselmis sp. 12.6+5.0 - 945 6600 7.0 0.0011 -.15 Total 38.6 2245 14600 21.4 0.0015 0.+ 3.5 4 9 Eutreptiella sp. and 11.7+3.7 75 5265 1400 3.3 0.0024 +.69 B M . lutheri and 11.0+6.0 - 550 8000 3.1 0.0004 -.10 e Isoselmis sp. 3.9+3.0 293 6600 1.1 0.0002 -.46 Total 26.6 6108 16000 7.5 0.0005 0 + 2.7 Stenosomella 9 Eutreptiella ap. 5.1+3.7 72 2295 1400 1.9 0.0013 ventricosa 0 + 1.8 0 + 3.5 12 M . lutheri 36.4+7.6 - 1820 8000 20.2 0J3025 +.15 and Ieoeelmia sp. 12.8+4.9 - 960 6600 7.1 0.0011 -.27 Total 49.2 2780 14600 27.3 0.0019 Eutreptiella sp. 1.6+1.2 70 720 1400 0.46 0.0003 -.08 and M . lutheri 17.0+6.8 - . 850 8000 4.9 . 0.0006 +.18 and Isoselmis sp. 5.0+4.6 - 375 6600 1.4 0.0002 -.32 Total 23.6 1945 16000 6.76 0.0004 . O n e - w a y A N O V A F-value Significant 26.4 Yes (.01) Comparisons (ScheffS's Test) with a significant difference at .05 level. 1/6. 1/7. 1/10. 1/12, 2/6. 2/7, 2/10, 2/12, 3/6, 3/10, 4/6, 4/8, 4/10, 5/8, 5/10, 6/8, 6/9, 6/11. 7/8, 7/11, 8/10. 8/12, 9/10, 10/11, 11/12. 82 and Stenosomella ventricosa (see Section 4b), were presented with three prey types i n various combinations. I t i s almost c e r t a i n that steady-state feeding conditions were reached during t h i s experiment despite the f a i r l y high feeding rates estimated. E u t r e p t i e l l a sp. was presented s i n g l y at 1,400 c e l l s / m l ; and together with Monochrysis l u t h e r i at(8,000 cells/ml)and Isoselmis sp. ; (6,600 c e l l s / m l ) . Tintinnopsis subacuta accumulated more E u t r e p t i e l l a sp. when presented s i n g l y than i n the three-prey mixture, but not s i g n i f i c a n t l y more. There was also not quite a s i g n i f i c a n t d i f f e r e n c e i n the number of M. l u t h e r i accumulated by T_. subacuta i n the two or three-prey mixture, but s i g n i f i c a n t l y fewer Isoselmis sp. were accumulated i n the three-prey mixture than i n the two prey mixture. The e l e c t i v i t y index f o r Isoselmis sp. (crypto-phyceae) was negative i n bothceases. As i n previous Tables the e l e c t i v i t y index f o r T_. subacuta feeding on E u t r e p t i e l l a sp. i n a mixture of prey types was highly p o s i t i v e . The feeding rate on the l a t t e r prey was also r e l a t i v e l y high - about 6 E u t r e p t i e l l a s p / h r / t i n t i n n i d or equivalent to 0.0042 ml/hr/tin-t i n n i d (see General Discussion). As shown i n Table 16 Stenosomella ventricosa accumulated s i g n i f i c a n t l y l e s s E u t r e p t i e l l a than did T_. subacuta (about 1/3 as many) when i t was pre-sented s i n g l y to the t i n t i n n i d s . S_. ventricosa also had l e s s E u t r e p t i e l l a sp. (but not s i g n i f i c a n t l y l e s s ) i n the 3-prey mixture than i n the single-prey case, and about'7-fold fewer than did j C . subacuta. E u t r e p t i e l l a sp. i n the 3-prey mixture had a negative e l e c t i v i t y index i n S^ . ventricosa . In contrast, the number of M. l u t h e r i accumulated by \S_. ventricosa was larger (but not s i g n i f i c a n t l y so) than the number i n T_. subacuta i n both the 2-prey and 3-prey mixtures. More M. l u t h e r i were found ins i d e S^ . ventricosa i n the 2-prey than i n the 3-prey mixtures, but again, the d i f f e r e n c e was not s i g n i f i c a n t . S^. ventricosa accumulated the same number of Isoselmis sp. as did T_. subacuta 83 i n the two-prey mixture, and a n o n - s i g n i f i c a n t l y larger number than did T. subacuta i n the 3-prey mixture. The e l e c t i v i t y index f o r S_. ventricosa on Isoselmis sp. was also negative i n both cases. In summary, these two species of t i n t i n n i d i n Table 16 responded i n a s i m i l a r manner to M. l u t h e r i and Isoselmis sp., and very d i f f e r e n t l y to the much larger E u t r e p t i e l l a sp.. I t i s i n t e r e s t i n g that the t o t a l number (but not the volume) of accumulated c e l l s i s s i m i l a r i n the 3-prey s i t u a t i o n for both species of t i n t i n n i d . In summary the r e s u l t s shown i n Tables 11 to 16 in d i c a t e various complex forms of d i f f e r e n t i a l predation f o r some t i n t i n n i d species on c e r t a i n prey types. In some cases t h i s behaviour could be said to be some form of negative s e l e c t i o n i n mixed-prey s i t u a t i o n s $ although i t s behavioural basis and adaptive value seem obscure. For example, Tintinnopsis parvula showed only d i f f e r e n t i a l predation i n favour of Monochrysis l u t h e r i over Isoselmis sp. i n Table 11 since the number of the l a t t e r accumulated was the same i n the single-prey as i n the two-prey s i t u a t i o n . However, the accumulation of M. l u t h e r i was not enhanced because i t was larger or f a s t e r than Isoselmis sp., as the reverse i s true. In Table 12 d i f f e r e n t i a l predation i s apparent by T_. subacuta feeding on E u t r e p t i e l l a sp. over Isochrysis galbana i n s i n g l e -prey s i t u a t i o n s , and negative s e l e c t i o n i s apparent against I_. galbana i n the 2-prey s i t u a t i o n . I t should be remembered that i n long experiments (Table 12) when a feeding steady-state has beemjrreached, that both apparent d i f f e r e n t i a l predation and negative s e l e c t i o n may i n fac t be caused by differences i n the ease with which the prey types are digested. In another long experiment (Table 13) D u n a l i e l l a t e r t i o l e c t a was accum-ulated l e s s i n a mixture with I_. galbana than when presented t e l . subacuta alone. The l a t t e r cannot be considered as either d i f f e r e n t i a l predation or 84 negative accumulation/selection since both prey types were accumulated i n proportion to t h e i r abundance. In a much shorter experiment (Table 14) T. subacuta and Tintinnopsis mucicola showed opposite d i f f e r e n t i a l predation; T_. subacuta aginst Isoselmis sp. and T. mucicola against E u t r e p t i e l l a sp.. In another short experiment (Table 15) T_. subacuta showed d i f f e r e n t i a l pre-dation against p_. t e r t i o l e c t a (as i n Table 13) when fed with M. l u t h e r i . This cannot be a product of the r e l a t i v e l y f a s t e r d i g e s t i o n of D. t e r t i o l e c t a , as the l a t t e r i s notably more d i f f i c u l t to digest by t i n t i n n i d s than i s M. l u t h e r i . As seen i n Table 15 the small t i n t i n n i d s T. rapa and T. nana show the maximum d i f f e r e n t i a l predation against D. t e r t i o l e c t a , almost c e r t a i n l y because i t i s too large for them. Stenosomella ventricosa shows d i f f e r e n t i a l predation on E u t r e p t i e l l a sp. when compared with T. subacuta (Table 16); and both t i n t i n n i d species show negative s s e l e c t i o n against Isoselmis sp. and M. l u t h e r i (to a l e s s e r extent) i n a 3-prey s i t u a t i o n . To gudge from the f i g u r e s i n Table 16 i t i s u n l i k e l y that the l a t t e r phenomenon i s caused by the fact that the feeding rates were only then at or above t h e i r maxima. E f f e c t s of p r i o r s t a r v a t i o n on feeding rates Tables 17 and 18 show the r e s u l t s of the e f f e c t of p r i o r starvation for various long periods on the feeding r a t e of Tintinnopsis subacuta and Stenosomella ventricosa. In the experiment shown i n Table 17, a dense but unknown concentration (but c e r t a i n l y above'the OFC l e v e l ) of D u n a l i e l l a  t e r t i o l e c t a was presented to T\ subacuta a f t e r the l a t t e r had been starved i n f i l t e r e d water for 0,30 or 48 hours. The non-starved c e l l s accumulated about twice as many food c e l l s as the starved c e l l s which a l l showed about the same r e s u l t s . These differences were not s t a t i s t i c a l l y s i g n i f i c a n t because of very large variance i n the samples. S i m i l a r l y i n Table 18,*tintinnids starved T A B L E 17. Tintinnopsis subacuta (etc.) starved for various periods in filtered seawater feeding on Dunaliella tertiolecta at unknown, but dense, concentrations. Salinity 26%,  Duration of Duration of Tempera- Number Number Number of other prior starva- feeding ture predators prey/ food items/ t i o n (hrs) p e r i o d (hrs) examined p r e d a t o r s T i n . Comments 30 48 nil 48 0.33 0.50 1.3 1.4 8 8 17 17 5 6 5 6 7.2+8.3 8.0+8.8 17.0+8.7 5.0+6.2 Tintinnidium mucicola nil nil 3.2 nil 3 had no food nil 1.3 17 nil 4.6 One-way ANOVA F - value Significant 2.98 no (.05) T A B L E 18. Tintinnopsis subacuta and other predators starved for various periods and feeding on Eutreptiella sp. Temperature 8 ° C ; Salinity 26&. Duration of Duration of prior starva- feeding tion ( h x a ) p e r i o d ( h r 3 ) Predator 72 72 72 90 90 90 90 0-4 © 0.4 1.0 © 0.45 © 0.45 © 1.1 © 1.1 © T . subacuta Synchaeta littoralis T . subacuta T . subacuta Stenosomella ventricosa T . subacuta S. ventricosa Number Number predators prey/ examined predator F R F R Number nos /hr / m l / h r / prey /ml predator predator Comments 3 5.0+1.0 14600 12.5 0.0009 Pred . cells thin; no storage granules 1 21.0 14600 52.5 0.0036 Pred . 400x200um 4 4.5+3.7 . 14600 4.5 0.0003 No granules 5 3.2+1.1 14600 7.1 0.00046 Pred . cells thin; no granules 3 1.3+1.2 14600 2.9 0.0002 Cel ls thin; some granules 5 8.4+1.2 14600 7.6 0.00051 No granules 3 ,4.3+1.6 14600 . 3.9 0.00026 Many granules One -way A N O V A F-value Significant Comparisons (Scheffg's Test) with a significant difference at .0 5 level 3.72 yes (.05) 4 / 5 87 fo r 72 or 90 hours i n f i l t e r e d water were then presented with very high concentrations of E u t r e p t i e l l a sp. (14,600/ml) for 0.4 or 1.0 hours. Both Tintinnopsis subacuta and Stenosomella ventricosa showed very low feeding rates on t h i s food whether starved f o r 72 or 90 hours, with T_. subacuta accumulating more E u t r e p t i e l l a sp. than S_. ventricosa (as i n Table 16). One i n d i v i d u a l of the large r o t i f e r Synchaeta l i t t o r a l i s contained about 4 times as many food items as did T_. subacuta (and see Section 4c) . Starved S^ . ventricosa contained some food storage granules although the c e l l s were t h i n , but the c e l l s of starved T_. subacuta were t h i n and without obvious food r e -serves. S_. ventricosa may be better adapted than T.' subacuta to environments containing l i t t l e food. The r e s u l t s of Tables 17 and 18 i n d i c a t e that s t a r v a t i o n for more than 48 hours can se r i o u s l y reduce the a b i l i t y of t i n -t i n n i d s to eat when food i s once more provided i n dense concentrations. Since the t o t a l number of c e l l s accumulated was small, i t i s not only a question of an i n a b i l i t y to digest food once eaten, because of low l e v e l s of the pools of d i g e s t i v e enzymes (and see discussion at the end of t h i s Section). Feeding and l o s s rates The experiments whose r e s u l t s are shown i n Tables 19 to 23 include estimations of the rate of 'loss' of old food items by t i n t i n n i d s , as well as estimations of feeding rates on newly presented prey'items. Losses are probably the r e s u l t of d i g e s t i o n plus egestion with the rate of digestion or p a r t i a l d i g e s t i o n as the l i m i t i n g step. Most food items contain an undiges-t i b l e f r a c t i o n , but th i s f r a c t i o n may very i n s i z e with changes i n the physio-l o g i c a l states of predator and prey. Therefore, i n Tables 19 to 23, l o s s rates r e f e r to the estimated rate of disappearance bf accumulated food items. It was not possible to know whether the food was a) mostly digested and then egested at a s i m i l a r r a t e ; b) wholly-digested with no residue to egest; or 88 c) egested before any d i g e s t i o n took place. As with feeding rates, losses are compared from differences between the mean values of s a c r i f i c e d sub-samples . In Tables 19 and 20 the r e s u l t s of a type of n u t r i t i o n a l 'homeostasis' i n t i n t i n n i d s are shown. When a natural seawater sample i s d i l u t e d with f i l t e r e d seawater of the same o r i g i n , there i s often very l i t t l e change i n the accumulated food c e l l s per t i n t i n n i d . This can r e s u l t from two possible causes: a) the d i l u t i o n of a dense concentration of p a r t i c u l a t e matter merely reduces that concentration to (or above) the OFC or l e v e l at which the t i n t i n n i d can feed (or digest) at i t s f a s t e s t r a t e ; or b) the l o s s rate of food i s reduced by the t i n t i n n i d to match the reduced opportunities for feed-ing i n a d i l u t e medium, thus increasing d i g e s t i v e e f f i c i e n c y and reducing the l i k e l i h o o d of starvation. The l a t t e r may be a l a r g e l y 'mechanical' con-sequence of a slower feeding r a t e . This problem i s also discussed l a t e r i n t h i s Section, In Table 19 i t can be seen that the small t i n t i n n i d species Stenosomella  n i v a l i s l o s t about h a l f of i t s accumulated food c e l l s i n 7 to 8 hours when a natural sample was d i l u t e d with f i l t e r e d seawater, whether the d i l u t i o n was 1/3 or 9/10 of the o r i g i n a l volume. Likewise, i n Table 20, the number of accumulated natural food items of Tintinnidium mucicola showed an appreciable but not s t a t i s t i c a l l y s i g n i f i c a n t d i f f e r e n c e a f t e r 5.5 to 7.3 hours, but only when the d i l u t i o n f actor was as great as 7 to 1. In d i l u t i o n s of 1 to 1 and 3 to 1 with f i l t e r e d seawater no d i f f e r e n c e was seen from the o r i g i n a l sample; and i n a l l three l e v e l s of d i l u t i o n , the change i n the accumulated c e l l number over about 24 hours p a r a l l e l e d the change (a s l i g h t decline) i n the undiluted o r i g i n a l sample. After 26 hours the T_. mucicola i n the. sample which had been T A B L E 19. Loos rate of Stenoaomella nivalis at two levels of dilution of medium with filtered seawater. Temperature 9°C; Salinity 26&. P e r c e n t a g e N u m b e r D u r a t i o n o r i g i n a l p r e d a t o r s ( h r B) s e a w a t e r e x a m i n e d Prey Number Percentage T i n B with Loss rate Loos rate prey/ proy algal n o s / T i n / u m 3 / T i n / predator digested evespots hr hr Condition 7.25 8.0 100 < 1 66 12 17 © ^5um g r e e n f l a g e l l a t e 8 21.1+11.3 10.2+8.1 8.0+10.5 51 77 7 8 1.5 1.6 ~75 - 8 0 Active Active Active One-way A N O V A T-value Significant Comparisons (Scheffe,'s Test) with a significant difference at .0 5 level 7.54 Yes (.01) 1/2, 1/3 co I T A B L E 20. Change of food contents of Tintinnidium mucicola with time at four levels of dilution of medium with filtered seawater. Temperature 9 ° C ; Salinity 25&. Time since Percentage Number dilution original predators ( h r s ) medium examined Prey Numbers Percentage prey/ prey predator dige sted 'Loss rate' f rom previous check nos /hr / Tin Nos & size of T i n . u m 3 / h r / storage Tin granules Tina dividing 5.5 24.0 6.5 24.5 7.0 24.7 7.25 26.5 100 100 50 50 25 25 12 12 6 10 7 9 10 5 Yellow-brown 22.3+13.8 cells (cryptos?) 15.7+7.6 " 24.1+9.8 •» 12.3+6'.6 " 26.7+11.3 " , 13.7+5.6 " 14.6+9.6 9.5+4.2 17 27 35 28 30 24 18 32 0.35 * ni l 0.66 * ni l 0.73 1.1 0.26 Many ->3um Many ->3um Many nil 1 ni l nil ni l 2 Few-low prey 1 number; Many-high number Very few nil Value for 100% medium at 5.5 hrs used ao "time sero" for this calculatic One-way A N O V A F-value 1 Significant Comparisons (Scheffe^s Test) with a significant difference at .05 level 3.31 yes (.01) ni l d i l u t e d by 7 to 1 had very few food reserve storage granules. T i n t i n n i d s i n the o r i g i n a l sample and i n 1 to 1 and'3 to 1 d i l u t i o n s s t i l l had many storage granules. This would seem to i n d i c a t e that there was s t i l l enough food to allow T_. mucicola to feed and store nutrients i n a l l samples, except i n that of the 7 to 1 d i l u t i o n . This experiment was c a r r i e d out at 9 C. T i n t i n n i d d i g e s t i o n (loss) rates may be f a s t e r at higher temperatures. Simultaneous feeding and l o s s rates Tables 21, 22 and 23 show the r e s u l t s of experiments i n which are e s t i -mated the rates of feeding on new food types and the simultaneous rate of l o s s of d i f f e r e n t o l d food types. Feeding and l o s s rates were estimated from the differences between the average number of food c e l l s / t i n t i n n i d , i n various samples since the s t a r t of the experiment. E f f e c t of temperature In Table 21 four t i n t i n n i d species Tintinnopsis subacuta, T. parvula; T_. rapa and Tintinnidium mucicola were presented with Monochrysis l u t h e r i at 5,000 c e l l s / m l and Cryptomonas minuta at 2,000 c e l l s / m l , whilst l o s i n g old natural food of various types (mostly unidentified) i n an experiment l a s t i n g up to 2 1/4 hours. The t i n t i n n i d s were taken from a f i e l d sample at 14.5 C and kept at one of the 4 experimental temperatures (8, 13, 17 or 22°C) f o r 6 hours before the experiment. One time check was made of those samples kept at 8 and 13°C and three time checks were made of samples at 17 and'22°C. Comparisons between l e v e l s (Scheffe's t e s t ) were confined w i t h i n the r e s u l t s for one species, to avoid undue complexity. In Table 21 the feeding rates of T_. subacuta on M. l u t h e r i are f a i r l y high and not s i g n i f i c a n t l y d i f f e r e n t at a l l 4 temperatures, with the maximum at 10.1 7 c e l l s / h r / t i n t i n n i d or equivalent to 0.0022 m l / h r / t i n t i n n i d . This 92 T A B L E 21. Tintinnopsis subacuta. T . parvula. T . rapa and Tintinnidium mucicola feeding on new food Monochrysis lutheri ( 42um 3 ) and Cryptomonas sp. (280am-1), and loss rate of old food of various types at four temperatures. Salinity 23 ,^,. Number Duration predators (hrs) Predator examined Numbers Volume Number F R or Temp. prey/ prey/ prey/ Loss-Rate F R a s predator predator m l n o s / h r / T i n m l / h r / T i n Elect iv i ty Index 0 + 2.0 T . subacuta © ® 8.0 0+1.5 T . subacuta © 10 13.0 Old 2.4 New: M . L . 17.4+10.0 New: Cryp . 0.8+1.8 Old: 1.5+1.4 New: M . L . 16.1+6. 6 New: Cryp . 0.7+0.8 731 224 676 197 5000 2000 5000 2000 8.7 0 .4 10.7 0 .5 0 .0017 0 .0002 0 .0022 0 .0002 +.15 - . 7 3 +.15 - . 7 4 T . subacuta © 0 + 1 . 6 T . subacuta ® © 17.0 Old: 17.0 6.2+1.1 Old: 3.8+2.7 New: M . L . 7.9+5.5 New: Cryp. ni l 332 n i l 5000 2000 1 . 6 4 . 9 n i l 0 .0010 n i l +.17 -1.0 0 + 2 .25 T . subacuta <8> 17.0 Old: 3.2+2.0 New: M . L . 5.6+1.7 New: Cryp. 0.2+0.4 235 56 5000 2000 F r o m 0 1.33 F r o m 0 2 .5 n i l 0 . 0 0 0 5 n i l +.15 -.81 T . eubacuta 22.0 Old: 10.1+4.3 0 + 1.0 T . subacuta 22.0 Old: 4.3+1.7 New: M . L . 4.2+4.4 New: Cryp. 0.4+0.4 5.8 176 5000 ' 4 . 2 112 2000 n i l 0 .0008 n i l +.12 -.54 0 + 1.5 T . subacuta 3 U~7> 22.0 O l d : 5.7+0.7 New: M . L . 8.0+_6.9 New: C r y p . 1.0 + 1.0 336 281 5000 2000 F r o m 0 2 .9 F r o m 0 5.3 0 .7 0 .0011 0.0004 +.11 .44 One -Way A N O V A F - v a l u c Si f . i i f icant 14.52 Yea (.01) C o m p a r i s o n s (Scheffe ' s Tent) with a s i g n i f i c a n t d i f f c r c n c <: at .0 5 l e v e l .  2/3, 2/4, 2/6, 2/12. 2/16. 3/5. 3/13. 4/13, 5/6, 5/12, 5/15. 5/16. 5719. 6/9. 6/13, 7/16. 9/12. 9/16, 12/13, 13/16 -. 93 T A B L E 21. Continued Duration (hrs) Predator Number predators Temp. examined °C 0 + 2.0 T. parvula © © © 0+1.5 T. parvula 0 T. parvula © 0 +1.6 T. parvula © ® 0+2.25 T. parvula ® 0 T. parvula 0+1.0 T. parvula © © 0+1.5 T. parvula © ® © Number Volume FR or prey/ prey/ Number Loss Rate FR as Electivity predator predator prey/ml nos/hr/Tin ml/hr/Tin Index 8.0 Old: 0.3+0.8 New: M.L. 14.0+6.1 New-. CM. 1.6+2.2 13.0 Old: nil New: M.L. 19.0+4.6 New: CM. 0.6+0.5 17.0 Old: -2.7+2.4 17.0 17.0 22.0 22.0 22.0 Old: 0.6+1.1 New; M.L. 12.3+3.3 N e w T c.M. nil Old: nil N e w : M.L. 14.6+5.6 N e w : CM. nil Old: 4.3+2.0 Old: 1.7+0.6 New: M.L. 4.3+5.1 New: CM. nil Old: 0.3+0.5 New: M.L. 14.0+6.7 New: CM. 0.3+0.5 588 450 798 169 517 nil 613 nil 181 nil 588 84 5000 2000 5000 2000 5000 2000 5000 2000 . 5000 2000 5000 2000 7.0 0.8 12.7 0.4 1.3 7.7 nU From 0 1.2 6.5 nil 2.6 4.3 nil F r o m 0 2.7 9.3 0.2 0.0014 0.0004 0.0025 0.0002 0.0015 nil 0.0013 nil 0.0009 nil 0.0019 0.0001 +.11 -.47 +.15 -.47 +.17 -1.-0 +.17 -1.0 + .17 -1.0 +.16 -.86 One-way ANOVA F-value S i gn i f i cant 27.22 yes (.01) C o m p a r i s o n s (Schcfff's Test) with a sign i f i c a n t difference at .05 level •  1/2. 1/4, 1/8. 1/9. 1/14. 2/3. 2/5. 2/6. 2/7, 2/13. 2/15. 3/4. 3/8. 3/9. 3/14. 4/5. 4/6. 4/7, 4/10. 4/11. 4/15. 5/8. 5/9. 5/14. 6/9. 7/8. 7/9, 7/14, 8/15. 9/13, 9/15, 13/14, 14/15. 94 T A B L E 2 1 . Continued Duration (hrs) Predator Number predators examined Temp. ° C Number prey/ predator Volume prey/ predator Number prey/ml FR or Loss Rate nos/hr /Tin. FR as ml/hr/Tin Electivity Index 0 + 2.0 T. mucicola © © 4 8.0 Old: 7.8+6.3 New; M.L. 0.5+0.6 New: CM. 0.8+1.0 21 225 5000 2000 0.25 0.40 0.00005 0.0002 -.30 +.37 0 +1.5 T. mucicola © © 5 13.0 Old: 11.8+5.8 New: M.L. nil New; CM. 0.6+0.6 nil 169 5000 2000 nil 0.40 nil 0.0002 - r . o + .56 0 T. mucicola 11 17.0 Old: -9.8+3.7 • _ — " 0 + 1.6 T. mucicola © © 8 17.0 Old: 8.0+3.6 New: M.L. 0.5+1.1 New: CM. 1.3+1.2 21 365 5000 2000 1.1 0.31 0.81 0.00006 0.0004 -.44 +.01 0 + 2.25 T. mucicola © . © 4 17.0 Old: 8.8+7.6 New: M.L. nil New: CM. 1.8+1.3 nil 506 5000 2000 . From 0 0.44 nil F rom 0 0.8 nU 0.0004 - 1 . 0 +.56 • 0 T. mucicola © . 6 22.0 Old: 7.7+2.6 - - _ _ .0 + 1J0 T. mucicola 0 •0 © 2 22.0 Old: 3.0+4.2 New: M.L. 2.0+2.0 New: CM. 1.0+1.0 84 281 5000 2000 4.7 2.0 1.0 0.0004 0.0005 -.04 +.07 0+1 .5 T, mucicola © © © 3 Old: 2.7+2.5 New: M.L. 1.3+2.3 New; CM. 1.0+1.7 55 281 5000 2000 From 0 3.3 From 0 0.86 From 0 0.66 0.00017 0.00033 -.12 +.21 One-way A N O V A F-value Significant Comparisons (Scheffc' difference at .05 leve l 6 Test) with a significant % 12.48 yes (.01) 2/4. 2/6. 2/7. 3/6. 4/5. 4/8, 4/9. 5/6, 5/7, 5/12, 6/8. 6/9, 7/8, 8/10, 8/12. T A B L E 21. Continued 95 Number Number Volume FR or Duration predators Temp. prey/ prey/ Number Loss rate FR as (hrs) Predator examined °C predator predator prey/ml nos/hr/Tin ml/hr/T Electivity in Indix 0 + 2.0 T. rapa 0+1.5 T. rapa 0 T. rapa 5 0 + 1.6 T. rapa 3 One-way ANOVA F-value Significant 3.21 No (.0 5) 8.0 13.0 17.0 Old: 3.0+0.8 New; nil Old: nil New. M.L. 1.7+1.7 New: C M . nil Old: 3.4+1.7 Old: 1.0+1.0 New; M.L. -0.7+0.8 New: C M . nil 71 29 5000 2000 5000 2000 1.0 0.0002 1.5 0.4 0.00008 Comparisons (Scheffg's Test) with a significant difference at .0 5 level.  nil + .17 -1.0 + .17 -1.0 0 + 2.0 Helicostomella 3 kiliensis 0+1.6 Helicostomella kiliensis 0+2.25 Helicostomella kiliensis 0 Helicostomella 1 kiliensis 0+1.5 Helicostomella 1 kiliensis 0 + 2.0 Stenosomella ventricosa 0 + 2.25 Stenosomella ventricosa 8.0 17.0 22.0 22.0 8.0 17.0 Old: 0.6+1.0 New: M.L. 13.3+5.7 New: C M . nil Old: nil New: M.L. 5.0+5.0 17.0 Old; nil . New: M.L. 6.0+6.0 Old: 7.0 Old: nil New; M.L. 9.0 New; C M . 2.0 New; M.L. 12.0 Old: nil New: M.L. 15.0 New: C M . nil 559 nil 210 252 378 562 504 630 5000 2000 5000 5000 5000 2000 5000 5000 2000 6.7 nU 3.1 2.7 4.7 6.0 1.3 6.0 6.7 0.0013 nil 0.0006 0.0005 0.0012 0.0007 0.0012 0.0013 + .17 -1.0 +.07 -.22 + .17 -1.0 96 o maximum was achieved at 13 C - the c l o s e s t temperature of those used to that of the f i e l d sample. The greatest feeding r a t e on M. l u t h e r i was at the f i r s t check (0 + 1.6 hours) at 17°C, but at the second check (0 + 1.5 hours) at 22°C. The number of C_'. minuta accumulated was much l e s s than that of M. o l u t h e r i at a l l temperatures and n i l at one check at 17 C. The e l e c t i v i t y value f o r T_. subacuta on (J. minuta was strongly negative at a l l temperatures again i n d i c a t i n g d i f f e r e n t i a l (negative) predation of t h i s t i n t i n n i d species on laboratory cultures of some cryptophyceae. The estimated r a t e of l o s s of o l d food from T_. subacuta tended to decline with time at both 17 and 22 C (and see discussion l a t e r i n t h i s Section). Loss rates could not be estimated at 8 and 13 C. The l o s s rate was greater at 22 than at 17 C; but again due to the small sample sizes and great i n d i v i d u a l v a r i a b i l i t y , there were no s i g n i f i c a n t d i f f e r e n c e s between the remaining numbers of old food c e l l s . In terms of biomass T_. subacuta d e f i n i t e l y l o s t a greater volume of o l d food than the other t i n t i n n i d species shown i n Table 21. Several of the T_. 3 subacuta contained one or two T_. nana (volume 3,000 ;um ) and E u t r e p t i e l l a sp. 3 (500 jum ) at the s t a r t , whereas the l a r g e s t old food c e l l s contained by T_. 3 parvula and T_. mucicola i n t h i s experiment were about 150 jum or l e s s i n volume. The feeding rate of Tintinnopsis parvula on Monochrysis l u t h e r i (Table 21) showed no s i g n i f i c a n t d i f f e r e n c e s at d i f f e r e n t temperatures or times, and the feeding rates s u r p r i s i n g l y , were s i m i l a r to those of the larger T_. subacuta (Table 21) at 8 and 13°C and greater than those of the l a t t e r at 17 and 22 C. As i n T_. subacuta the estimated feeding r a t e by T. parvula on M. l u t h e r i rose to i t s maximum e a r l i e r at 17 C than at 22°C. Since 22°C i s much higher than the highest f i e l d temperature experienced by these species, t h i s 'lag' i n the feeding rate at 22°C may r e f l e c t some problem of 97 p h y s i o l o g i c a l adaptation, but i f so i t does not appear i n the differences between loss rates. The feeding rates of T_. parvula on Cr yptomonas minuta were, l i k e those of T_. subacuta, very low or n i l and the e l e c t i v i t y values were always negative. The accumulated number of M. l u t h e r i c e l l s i n Tintinnidium mucicola (Table 21) was very much lower than i n T_. subacuta and' T. parvula despite a comparable contact r a t e . The number of C. minuta per T_. mucicola was very small at a l l temperatures and only s l i g h t l y greater than that i n T_. subacuta . and T. parvula. However, the e l e c t i v i t y index of T_. mucicola on C. minuta was p o s i t i v e , unlike that for the other two t i n t i n n i d species. This r e s u l t emphasizes a problem i n the use of Ivlev's index, i n that i t s magnitude and even i t s sign depends not only on the amount eaten of the food item i n question, but also on the amount of the other items eaten. The l o s s rate o of o l d food from T. mucicola at 22 C was higher than that of T_. parvula and lower than that of T. subacuta. The l o s s rate i n Table 21 was also higher at 22 C than at 17°C, and declined with time (or more l i k e l y with biomass remaining) as d i d the l o s s rates of T_. parvula and T_. subacuta. Fragmentary r e s u l t s from a few other species of t i n t i n n i d i n the same experiment as above are shown i n Table 21. Tintinnopsis rapa contained very l i t t l e of either new prey species at three of the experimental temperatures, but i t s rates of l o s s of o l d food material at 17 C were no lower than those of the larger species. The sample sizes of Helicostomella k i l i e n s i s were extremely small, but i f the information can be u t i l i s e d , then t h i s species seemed to contain about as many c e l l s of M. l u t h e r i at 8 and 17 C as d i d T_. subacuta and T. parvula. The r a t e of l o s s of o l d food material by H. k i l i e n s i s at 22°C was also i n the same range as that of J_. subacuta and T_. parvula. 98 One c e l l of Stenosomella ventricosa i n each sample at 8 and 17°C, also con-tained a number of M. l u t h e r i c e l l s i n the same range as T_. subacuta and T_. parvula, and no Cryptomonas sp.. The incidence of the early stages of r e -production i n t h i s experiment was rather higher i n Tintinnidium mucicola than i n T_. subacuta or T_. parvula but was apparently not p o s i t i v e l y correlated with higher temperatures. E f f e c t of Starvation Table 22 shows the r e s u l t s of an experiment i n which Tintinnopsis  subacuta was presented with a dense mixture of Monochrysis l u t h e r i (16,000/ml) and Plagioselmis sp. (8,000/ml) for 3 hours, and checks were made of the number of accumulated food c e l l s / t i n t i n n i d at 0 + 1 . 0 and at 0 + 3.0 hours. At 0 + 2.17 hours some of the l a t t e r T_. subacuta were gently washed with a large volume of f i l t e r e d water, and t i n t i n n i d s i n one sub-sample of these were starved i n f i l t e r e d water f o r 0.66 hours. A second sub-sample of washed T_. subacuta was presented with a mixture of two new prey types: E u t r e p t i e l l a  sp. (3,800/ml) and Isoselmis sp. (15,000/ml) f o r 0.33 hours. T_. subacuta accumulated both species of o l d food i n the o r i g i n a l mixture i n the propor- -tions i n which they were presented, but rather slowly. There were no s i g -n i f i c a n t d ifferences between the numbers of M. l u t h e r i , nor between the numbers of Plagioselmis sp. per t i n t i n n i d i n any of the treatments. However, i f the differences between the accumulations of o l d food are used to estimate l o s s rates on an hourly b a s i s , i t i s highly probable that the l o s s rate of o l d food ( e s p e c i a l l y M. l u t h e r i ) was greater i n the T_. subacuta fed with new food than i n those c e l l s which ewere starved. The hourly los s r a t e of M. l u t h e r i was 4.1 c e l l s i n the starved c e l l s a f t e r 0.66 hours and 22.8 c e l l s i n the newly fed T_. subacuta (after 0.33 hours). This comparison between samples taken at d i f f e r e n t times i s also p a r t l y specious because l o s s rates T A B L E 22. Feeding and loss rates of Tintinnopsis subacuta; losing Monochrysis lutheri and Plagioeelmls ap. and either starved, or gaining Eutreptiella sp. and Isoselmis sp. Number Number F R or F R or L R L R Gain (D) Duration predators _ prey/ Number Loss Rate Loss Rate F R D - C D - C Loss (D-C) (hrs ) e x a m i n e d Prey predator prey/ml n o s / h r / T i n u m 3 / h r / T i n m l / h r / T i n no3 / h r / T i n u m 3 / h r / T i n u m 3 A 1.0 21 © © M . lutheri a n d Plagioselmis sp. 16.1+6.2 8.0+3.2 16000 8000 16.1 8.0 80 5 600 0.0010 0.0010 - . - -B 3.0 18 © © M . lutheri a n d Plagioselmis 24.2+6.0 13.0+4.3 16000 8000 -_ • C 0.66 (starved from 2.17 hrs) 18 © © O l d : M . lutheri a n d Plagioselmis N e w Food 19.3+6.2 9.8+3.7 nil ni l ni l ni l 4.1* 3.3* nil 205 '396 - - -D 0.33 ( n e w food after 2.17 hrs) 7 © © © O l d : M . lutheri a n d • Plagioselmis N e w : Eutreptiella a n d Isoselmis 14.4+7.3 11.0+4.0 2.4+1.0 2.4+2.4 ' n i l ni l 3800 15000 22.8* 3.0* 7.2 7.2 1140 360 3744 540 0.0019 ' 0.00048 18.7 nil 935 nil 4.6 *Loaa rates calculated from interpolated values A to B of accumulated old food at 2.5 hours. O n e - w a y A N O V A F - v a l u e S i g n i f i c a n t 23.63 y e s (.01) Comparisons (Scheffe's Test) with a significant difference at .0 5 level. 1/2, 1/6. 1/9, 1/10, 2/3, 2/5, 2/9, 2/10. 3/6. 3/9. 3/10, 4/9, 4/10, 5/6, 5/9, 5/10, 6/10, 7/9. 7/10. 8/9. 8/10. 100 dec l i n e with time. However, a 'forcing e f f e c t ' of the ingestion of new food on the d i g e s t i o n (or disappearance) of old food has also been observed i n the c i l i a t e Stentor coeruleus (D.J. Rapport, unpublished data) and i s probably a r e a l phenomenon (but see Table 23). The accumulation of new food shown i n Table 22 was rather slow (0.0019 and 0.00048 ml/hr/tin) and was much less than the (estimated) l o s s r a t e of old food i n terms of numbers of c e l l s , but i n terms of biomass the r a t i o of hourly gain/loss e n t i r e l y due to feeding (D-C i n Table 22) was 4.6/1. This phenomenon,.which w i l l be discussed again l a t e r i n t h i s Section, was due l a r g e l y to the f a c t that one of the new prey types, E u t r e p t i e l l a sp. i s much larg e r than the others used i n t h i s experiment. Tables23 shows the r e s u l t s of a s i m i l a r experiment where three species of the 'warm-water'type of t i n t i n n i d s , namely Tintinnopsis c y l i n d r i c a , Helicostomella k i l i e n s i s and Eutintinnus latus plus the ubiquitous Tintinnidium  mucicola, were given food f o r several hours then washed i n f i l t e r e d water and eith e r starved, or a l t e r n a t i v e l y given new food of a v i s i b l y d i f f e r e n t type. The o ld food type used was one of either Monochrysis l u t h e r i , Isoselmis sp. or D u n a l i e l l a t e r t i o l e c t a ; and i f given new food, the type used was either Isoselmis sp. or p_. t e r t i o l e c t a . Scheffe's test of multiple comparisons was not made between the r e s u l t s from d i f f e r e n t t i n t i n n i d species. The r e s u l t s of t h i s experiment (Table 23) seem to be p a r t l y i n d i r e c t contrast to those shown for another species of t i n t i n n i d i n Table 22, i n that o l d food c e l l s of M. l u t h e r i or I), t e r t i o l e c t a were not l o s t more rap-i d l y from those T. c y l i n d r i c a c e l l s given new food than from those starved. However, i n the case of old Isoselmis sp. there were s i g n i f i c a n t l y more old food c e l l s remaining i n starved T_. c y l i n d r i c a than i n those fed with new T A B L E 23. Accumulation and loss rates of Tintinnopals cylindrica, HelicoatomeUa kiliensis, Tintinnidium mucicola and Eutintinnus latus, feeding on M o n o c h r y s i 3 lutheri, or Isoselmis sp.; or starved, and losing M . lutheri, Isoselmis 3 p . , o r Dunaliella tertiolecta. Temperature 1 6 ° C ; Salinity 16&. Duration (hrs) Predator Number predators examined Prey Number prey/ predator Percentage prey dice sted Number s prey /ml F R or L R n o s / h r / T i n FR or L R U m 3 / h r / T i n F R m l / h r / T i n 0 Tintinnopsis cylindrica 0 10 M . lutheri 53.9+12.2 39 ~ — 6.7 ii © 6 Old: M . L . 8.3+5.2 38 nil 6.8 462{L) -(starved) 7.17 II © 9 . Old: M . L . 8.6+9.4 75 ni l 6.3 430(L) -* and New Isoselmis ap. 3.9+3.9 26 13000 ,0.54 43(F) .0.00004 0 II © 6 D. tertiolecta 16.7+6.0 • 50 - - -7.17 II © 7 Old: D . T . 3.4+2.6 88 nil 1.85 615{L) -(starved) 7.66 II © 11 Old; D . T . 2.1+3.1 100 nil 1.91 635(L) -'• and New © M . lutheri 24.8+11.0 6 4000 3.24 220(F) 0.0008 0 it © 11 Isoselmis s p . 14.8+7.5 61 - - - -8.0 II 7 Old: Isoaelmis s p . 8.6+2.5 100 nil 0.78 62(L) ->—' (starved) 8.33 II © 5 Old Isoselmis s p . 1,0+1.4 100 nil 1.66 133(L) -and New M . lutheri 41.6+19.2 40 3500 5.00 340(F) 0.0014 One-way A N O V A F-value Significant 29.01 yes (.01)' Comparisons (Scheffe's Test) with a significant difference at .05 level 1/2, 1/3, 1/4, 1/6, 1/7, 1/9. 1/10. 1/11. 3/8, 3/11. 3/12, 4/5, 4/8. 4/9. 4/12. 5/6. 5/7, 5/11, 6/8. 6/9. 6/12. 7/8. 7/9. 7/10. 7/12. 8/11, 9/11, 10/11, 11/12. Duration (hrs) Predator Number predators examined • 6.7 0 7.66 0 8.33 0 7.17 7.66 0 8.0 8.33 Hclicostomclla kilienaia 2 2 1 2 2 One-way A N O V A F-value Significant 0 . 6 1 Tintinnidium mucicola 4 2 3 4 1 1 M . lutheri Old M . L . D. tertiolecta Old D . T , and New M . lutheri 8.0+7.0 2.0+1.4 5.5+2.1 0.0 . 0.0 Iaoaelmia sp. 6.0+1.4 Old Isoselmia sp. 0.0 and New M . lutheri 11.5+9.1 M . lutheri ' D . tertiolecta Old D . T . Old. D . T . and New M . lutheri Isoselmia sp. Old Isoaelmis sp. 4.0 Old Isoselmis sp, 0.0 and New M . lutheri 0.0 4.0+1.9 ni l ni l ni l ni l 18.3+13.9 \ Percentage ' prey Numbers FR or, LR FR or LR FR • digested p r e y / m l n o s / h r / T i n | i m 3 / h r / T i n m l / h r / T i n 12 - - - -50 n i l 0.9 45 -(starved) 0— 36 ni l ni l 0.72 239 ni l 4000 ni l n i l n i l 50 . . . nil n i l 0.72 58 74 3 500 1.38 110 0.00 0 39 60 nil - - -nil ni l — (starved) nil - - -nil - -40 - - -100 ni l 1.78 143 (starved) ni l ni l 2.20 176 ni l 3500 ni l ni l ni l TABLE 23. Continued Duration (hrs) Predator Number predatorB examined Prey Number Percentage prey/ prey Numbers FR or LR FR FR pr LR predator digested prey/ml nos/hr/Tin um 3/hr/Tin ml/hr/Tin 6.7 7.17 7.66 0 0 Eutintinnus latus E. tubulosus M. lutheri M. lutheri 10.0 8.0 Old D. tertiolectus 10.0 Old D. tertiolectus 2.0 and New. M. lutheri 6.0 Isoselmis sp. D. tertiolecta 8.0 8.0 50 33 100 100 100 88 33 nil (starved) nil (starved) 4000 0.3 0.78 20 261 0.0002 T A B L E 23A. Summary of accumulation experiments with T i n t i n n o p s i s subacuta. M a x i m u m Table Max. F R Numbers A v ; P r e y Max. P r e y D i f f l . Apparent Number P r e y m l / h r / T i n . p r e y / m l N o s / T i n . V o l u m e / T i n . P r e d . Selection 7 D. t e r t i o l e c t a 0.0065 1, 750 10 D. t e r t i o l e c t a 0.0018 15, 600 12 E u t r e p t i e l l a sp. 0.0004 4, 400 12 I. galbana 0.00004 66, 000 13 D. t e r t i o l e c t a 0.00034 4, 400 13 E u t r e p t i e l l a sp. 0.0018 1, 500 13 I. galbana 0.00011 66, 000 14 E u t r e p t i e l l a sp. 0.0023 3, 650 14 I s o s e l m i s sp. 0.00010 . 30, 000 1 5 M . l l u t h e r i 0.0020 13, 000 1 5 D. t e r t i o l e c t a 0.0003 6, 200 16 E u t r e p t i e l l a sp. 0.0042 1, 400 16 M. lu t h e r i 0.0018 8, 000 16 I s o s e l m i s sp. 0.0011 6, 600 21 M. lu t h e r i 0.0017 5, 000 21 Cryptomonas sp. 0.0004 2, 000 22 M. l u t h e r i 0.0010 16, 000 22 P l a g i o s e l m i s sp. 0.0010 8, 000 22 E u t r e p t i e l l a sp. 0.0019 3, 800 22 I s o s e l m i s sp. 0.00048 15, 000 32.5 6, 825 - --25.1 4, 718 18.3 9, 100 Y e s No 26.3 1, 200 Y e s Y e s 9.7 2, 040 Y e s Y e s 16.0 8, 000 Y e s No 46.7 2, 100 Y e s No 5.3 2, 650 Y e s No 2.0 150 Y e s No 26.5 1, 325 Y e s No 2.0 400 Y e s No 16.0 7, 200 Y e s Y e s 26.0 1, 300 Y e s Y e s 12.6 945 Y e s Y e s 17.4 730 Y e s No 1.0 280 Y e s No 24.2 No No 13.0 No No 2.4 Y e s No 2.4 Y e s No 104 M. l u t h e r i . This may be due to the f a c t that many more new food c e l l s were accumulated i n 7 to 8 hours by the T. c y l i n d r i c a i n d i v i d u a l s containing old Isoselmis sp.(and thus the smallest t o t a l volume of old food) than i n the other two cases. However, the d i f f e r e n c e between the accumulations of new M. l u t h e r i i n T_. c y l i n d r i c a containing o l d D. t e r t i o l e c t a or old Isoselmis sp. was not s i g n i f i c a n t . Hence, the 'forcing e f f e c t ' of new food on o l d food may depend somewhat on the amount of new food eaten. The small los s of old M. l u t h e r i during the very small accumulation of Isoselmis sp. seems to emphasize t h i s point. It i s i n t e r e s t i n g that T_. c y l i n d r i c a (as T_. subacuta and Stenosomella  ventricosa i n previous tables) shows d i f f e r e n t i a l (negative) predation/accum-u l a t i o n on Isoselmis sp. The r e l a t i o n s h i p of old and new food i s also d i s -cussed l a t e r i n t h i s Section. The gain/loss r a t i o f or T_. c y l i n d r i c a was p o s i t i v e i n terms of numbers when new M. l u t h e r i followed old 1). t e r t i o l e c t a or old Isoselmis;; but was p o s i t i v e i n terms of c e l l volumes only i n the l a t -ter case. This i s because M. l u t h e r i and Isoselmis sp. are of s i m i l a r s i z e 3 3 (50 and 75 /am ) but I), t e r t i o l e c t a i s much larger (200 jam ) . The sample sizes of Helicostomella k i l i e n s i s i n Table 23 were too small to show s i g n i f i c a n t differences between means even i f they had existed, but the rates of feeding and l o s s i n t h i s species were lower than i n the con-siderably l arger T_. c y l i n d r i c a , except that old Isoselmis sp. was l o s t at about the same rate i n the two species. Table 23 also shows again the apparent r e l a t i v e a f f i n i t y f o r Cryptomonad c e l l s by Tintinnidium mucicola. The accumulation and los s rates of t h i s t i n t i n n i d species on Isoselmis sp. were not only much larger than on the other two food types, but were also s l i g h t l y l a r g e r than the feeding or l o s s rates of T. c y l i n d r i c a or 105 H. k i l i e n s i s on Isoselmis sp. The l o s s r a t e of one i n d i v i d u a l of T_. mucicola given the opportunity to eat M. l u t h e r i (although i t d i d not do so) was great-er than that of one starved T. mucicola c e l l . It i s possible that r e l a t i v e ' a c t i v i t y ' (eg. 'handling'or avoiding p a r t i c l e s ) has some e f f e c t on the rate of l o s s of o l d food material. The l o s s rate of o l d J). t e r t i o l e c t a from one starved Eutintinnus latus was slower than i n one E. l a t u s c e l l fed with new M. l u t h e r i (Table 23). In summary, the r e s u l t s of Tables 21, 22, and 23 i n d i c a t e that feeding rates i n four species of t i n t i n n i d are l i t t l e affected by changes i n tem-perature a f t e r acclimation for several hours (Table 21); but that simultaneous l o s s rates are somewhat increased at very high temperatures?. Table 21 also showed once more the apparent d i f f e r e n t i a l predation of T. subacuta and T. parvula against a cryptomonad prey, and of T. mucicola against a non-crypto-monad prey. The l o s s rates ( i n numbers) varied d i r e c t l y with the c e l l s i z e of the t i n t i n n i d species, as d i d the feeding rates i n a general sense. Loss rates (in biomass) were greatest for T. subacuta, since some of i t s o l d food items were extremely large. In Tables 22 and 23 the old food items were of known species, and p a r a l l e l experiments were done with starved and fed t i n -t i n n i d s . T_. subacuta as shown i n Table 23, for once di d not eat proportion-' a t e l y l e s s of a cryptomonad (Plagioselmis sp.) then of M. l u t h e r i ; and when starved, T. subacuta also l o s t both of these prey types i n proportion. How-ever, negative d i f f e r e n t i a l predation was seen against the new cryptomonad food Isoselmis sp. when compared with the new E u t r e p t i e l l a sp. i n Table 22. In Table 22 i t can be seen that o l d food i s l o s t more r a p i d l y when new food i s added, than when the t i n t i n n i d i s starved. An a l t e r n a t i v e f(or additional) explanation f o r these r e s u l t s i f that l o s s rates decline exponentially with time whether the t i n t i n n i d i s fed or not. 106 Table 23 shows that the l o s s rates of old food from TiritirinOpsis  c y l i n d r i c a i n a long ( 7 - 8 hour) experiment were i n some cases a function of the amount of new food eaten (or vi c e - v e r s a ) , which i n turn depended upon the type of new food presented. T_. c y l i n d r i c a was the fourth species i n t h i s study to show negative d i f f e r e n t i a l predation on the cryptomonad Isoseihmis sp. Individual v a r i a b i l i t y i n t i n t i n n i d s l o s i n g o l d food and eating new food In the previous experimental r e s u l t s there appeared to be a p o s i t i v e r e -l a t i o n s h i p between the average amount of new food added to, and the average amount of old'food concurrently l o s t from t i n t i n n i d s . It was not cl e a r which of the two processes was the c o n t r o l l i n g factor i n the r e l a t i o n s h i p . I t i s probable that the process of ingestion of food i n some way 'forces' the t i n -t i n t i n n i d to 'lose' food more r a p i d l y ; and starving c e l l s or c e l l s with r e l a t i v e l y l i t t l e food seem to lose i t at a slower rate than do well-fed c e l l s (Table 22, 23; and Rapport, unpublished data). Also Berger (1971) and Goulder (1972) have shown that o l d food disappears from some other c i l i a t e s at a rate which i s much f a s t e r i n i t i a l l y than l a t e r . This i s also a well known phenomenon in« during the egestion of s o l i d p a r t i c l e s and the excretion of metabolites by planktonic Crustacea. The amount of o l d food i n s i d e a t i n t i n n i d may also place some r e c i p r o c a l upper l i m i t on i t s feeding rate on new food. There may be an upper l i m i t f o r the t o t a l number or volume of a l l contained food c e l l s which i s constant f o r a p a r t i c u l a r t i n t i n n i d species, and at which ingestion rate equals the maxi-mum d i g e s t i o n rate. If t h i s 'reservoir s i z e ' i s approximately a constant then the t o t a l of old and new food should be very s i m i l a r i n d i f f e r e n t i n d i -v i d u a l s i n one experiment, no matter what the v a r i a b i l i t y c o f t e i t h e r o l d or new 107 food per t i n t i n n i d . Also, i f t h i s theory holds the mean values of o l d and new food i n several t i n t i n n i d s i n each of a successive s e r i e s J of samples,, should l i e on a negatively sloping regression l i n e of old (x) on new (y) food; and the i n d i v i d u a l values should f a l l c lose to such a l i n e with r e l a -t i v e l y small variance. T h i r d l y to f u l f i l the predictions of t h i s theory, the values of the two intercepts extrapolated from a l i n e j o i n i n g the means of successive samples should be s i m i l a r and be equivalent to the mean t h e o r e t i c a l r e s e r v o i r s i z e for that species ( i e . the l i n e should have a slope of -1.0). As an i n d i c a t i o n that the 'constant r e s e r v o i r ' theory i s u n l i k e l y to be true, i t can be calculated from Tables 21 to 23 that there i s a wide range i n average numbers of old plus new food c e l l t o t a l s for each species i n various parts of those experiments. The range i s 8.9 to 30.2 t o t a l food i t e m s / t i n -t i n n i d f o r Tintinnopsis subacuta; 12.5 to 53.9 for T_. c y l i n d r i c a ; 6.0 to 19.6 for T_. parvula and 5.0. to 12.2 for Tintinnidium mucicola. The ranges of-., hourly gain to loss r a t i o s f o r averages of these species can also be c a l c u -l a t e d from Tables 21 to 23, and these too are f a i r l y wide. These ranges are: for Tintinnopsis subacuta 0.72 to 3.1 (number) and 2.9 to 4.6 (volume); f o r T_. c y l i n d r i c a 0.08 to 3.0 (number) and 0.1 to 2.0 (volume); for T_. parvula 1.7 to 3.5 (number) and for Tintinnidium mucicola 0.46 to 1.8 (number). The r e s u l t s of two experiments done 4 days apart to test t h i s theory are shown i n Figures 6 and 7. F i r s t l y i t i s obvious that the variance about any regression l i n e i n Figures 6 and 7 i s very large, and that the r e s e r v o i r theory mentioned above does not hold i n terms of volume. In f a c t , as the necessary conditions for normal l i n e a r regression analysis do not apply to Figures 6 and 7, a constrained l i n e a r regression analysis was performed. The l i n e was constrained about the Y axis at a point equivalent to the mean of 108 Figure 6. Relationship between the volume Q i m J x 10") of o l d food and new food contained by i n d i v i d u a l Tintinnopsis subacuta. new food - D u n a l i e l l a t e r t i o l e c t a o l d food - Isochrysis galbana and natural food items. O Low concentrations of I. galbana checked at 0 + 5 mins. • Low concentrations of I. galbana checked at 0 + 57 mxns. A High concentrations of I. galbana checked at 0 + 8 mxns. A High concentrations of I. galbana checked at 0 + 68 mins. 72r 68 64 60-56 52 48 VOLUME 4 4 of NEW FOOD 40 {.Jim3x 1 0 2 ) 3 6 32 ® 0 9 Y = - 0 . 8 5 ( X - 0 ) + - 2 6 5 0 R 2= 0 . 0 7 4 8 12 "16 2 0 V O L U M E o f O L D F O O D 24 2 8 ( p m 3 X 1 0 2 ) 110 3 2 Figure 7. Relationship between the volume (um x 10 ) of old food and new food contained by i n d i v i d u a l Tintinnopsis subacuta and Stenosomella ventricosa./ [-mean value O - T, Subacuta -©-.-— S_. ventricosa 0^^ ) new food - D u n a l i e l l a t e r t i o l e c t a old food - various O - T i n t i n n i d s i n f i l t e r e d water for 7 hours. • - T i n t i n n i d s from f i e l d sample. A - T i n t i n n i d s from f i e l d sample plus high concentrations of 5 u^m d i a . p o l y s t y r e n e l a t e x . A - T i n t i n n i d s from f i e l d sample plus high concentrations of Isochrysis galbana. 72 » 68 • 64 -60 * A 56 52 • 9 48 - • VOLUME 44 &• • of NEW 40 -A • FOOD 36 0 (Dunaliella 3 2 A A 9 Y = -1.9(X-0) + 3600 R2* 0.141 tertiolecta A ( p 3 X 1 0 2 ) 5 6 7 8 10 11 12 13 14 15 16 17 18 19 20 VOLUME of OLD FOOD ( j J 3 X 1 0 2 ) (Isochrysis galbana) 42 43 £ 112 a l l Y values when X (old food) was zero. The regression analysis shown i n Figure 7. was performed only on data from Tintinnopsis subacuta. In these circumstances, the regression of X on Y would n a t u r a l l y be s i g n i f i c a n t l y 2 d i f f e r e n t from zero. However, the regression c o e f f i c i e n t (R ) i n both Figures 6 and 7 i s very small, i n d i c a t i n g great i n d i v i d u a l v a r i a b i l i t y i n the data. The values (volume) of the X and Y intercepts are very s i m i l a r i n Figure 6 where the old food was D u n a l i e l l a t e r t i o l e c t a and the new food was Isochrysis galbana etc. However, the value of the X intercept i n Figure 7 i s about h a l f that of the Y intercept value, and here the o l d food i s either I. galbana, 5 yum l a t e x , or small natural p a r t i c l e s , a l l smaller than the new food D. t e r t i o l e c t a . Therefore the intercept values of X and Y i f based upon food c e l l numbers would not be s i m i l a r i n eit h e r experiment. I t i s curious that the mean values of the various sub-groups i n Figures 6 and 7 do f a l l close to an imaginary s t r a i g h t l i n e despite the large scatter of the i n d i v i d u a l values. A one-to-one c o r r e l a t i o n between the intercept volumes might be expected i f the l i m i t a t i o n on feeding rate was set i n some way by the t o t a l volume of food i n s i d e the t i n t i n n i d undergoing d i g e s t i o n . A one-to-one c o r r e l a t i o n between the intercepts i n terms of numbers of o l d and new food items might be expected i f each item i s contained i n a d i f f e r e n t food vacuole, as i s usually the case p a r t i c u l a r l y for large items, and i f the formation of food vacuoles i s the l i m i t i n g f a c t o r i n the process of ingestion. C e r t a i n l y food vacuole membrane synthesis i s an extremely active process i n some phagotro-phic protozoa (Ricketts, 1971); but recent work with the c i l i a t e Paramecium caudaturn (Allen, 1973) has indicated that the food vacuole membrane i n t h i s species i s neither f i n a l l y egested with the food residue nor broken down into i t s components, but i s 'conserved' as fragments i n small v e s i c l e s which 113 may be r a p i d l y transported back to the o r a l region for reuse. On the other hand, Ricketts (1973) states that the d i g e s t i v e enzymes within food vacuoles i n Tetrahymena pyriformis are not conserved at egestion, but are l o s t to the medium. Ricketts thinks that the ingestion rate of T_. pyriformis i s u l t i -mately l i m i t e d by the a v a i l a b i l i t y of d i g e s t i v e enzymes. An experiment with Paramecium a u r e l i a showed that food vacuoles containing old red carmine p a r t i c l e s were reduced i n number at the same rate as vacuoles containing new black ink p a r t i c l e s were formed by the c i l i a t e (J.D. Berger, unpublished data). If there i s any correspondence at a l l between the l o s s of old food and the gain of new food i n t i n t i n n i d s , i t i s perhaps a l i t t l e more l i k e l y e s p e c i a l l y for large prey to be on a number (or food vacuole?) basis than on the basis of r e l a t i v e volumes or biomass of material. This i s a subject about which very l i t t l e i s known f o r any protozoan. There are several other possible explanations for the great v a r i a b i l i t y i n feeding rates between i n d i v i d u a l t i n t i n n i d s seen i n the r e s u l t s of these accumulation experiments. These are that v a r i a b i l i t y may be due to one or more of the following: 1) Heterogenous d i s t r i b u t i o n of food items i n the environment; 2) the s i z e of the unused portion of the poolcfof d i g e s t i v e enzymes i n the c e l l , which i n turn may be the product of (a) the recent feeding rate and biomass of prey, and (b) the less-recent n u t r i t i o n a l h i s t o r y and the subsequent r a t e of synthesis of d i g e s t i v e enzymes; 3) d i f f e r e n t i a l p h y s i o l o g i c a l e f f e c t s of the immediate environment, other than food; 4) short-term spontaneous changes i n motion or feeding behaviour; 5) the age of the t i n t i n n i d c e l l or how recently the parent c e l l divided; 6) genetic v a r i a b i l i t y among members of a clone or between s t r a i n s or syngens within a species. Most of these p o s s i b i l i t i e s have not been tested i n t i n t i n n i d s , nor i n any protozoan. 114 (1) The heterogenous d i s t r i b u t i o n of food items may be a fa c t o r par-t i c u l a r l y i n l e s s dense c e l l concentrations or when experimental vessels are not agitated. In f a c t , experiments with very dense c e l l concentrations or with s t i r r e d samples seem to contain as much i n d i v i d u a l feeding v a r i a b i l i t y as any others. (2) D i f f e r e n t i a l d i g e s t i v e enzyme synthesis and u t i l i s a t i o n i s a l i k e l y p o s s i b i l i t y which could have complex e f f e c t s on the future feeding performance of a t i n t i n n i d . T h e o r e t i c a l l y , larger species should have a greater capacity f o r enzyme synthesis and storage than smaller species, and therefore should be l e s s i n d i v i d u a l l y v a r i a b l e ; and previously well-fed i n d i v i d u a l s of any species should be better able to recover from starvation when once re-fed, than i n d i v i d u a l s whose h i s t o r y has been one of poor nut-r i t i o n (and see Ricket t s , 1973). Feeding on 'blooms' or dense concentrations of the same species or q u a l i t y of prey type, should r a p i d l y reduce the variance i n feeding performance i n a t i n t i n n i d species caused by factor (2). Fenchel (1968) has shown that f o r many species of benthic c i l i a t e the maximum reproductive rate decreaseswwith increasing c e l l volume. As t h i s i s probably also true f o r t i n t i n n i d s i t i s l i k e l y that when a l l species are at (or near) t h e i r maximum reproductive rate, the v a r i a t i o n between c e l l s due to d i f f e r e n t n u t r i t i o n a l h i s t o r i e s w i l l be reduced more r a p i d l y i n small species than i n large ones. However, small species almost c e r t a i n l y need much denser con-centrations of food to reach t h e i r maximum reproductive rate than do large species (see General Discussion). (3) D i f f e r e n t i a l p h y s i o l o g i c a l e f f e c t s of unknown o r i g i n are also a d i s t i n c t l y p ossible source of v a r i a t i o n i n i n d i v i -dual feeding rates, and t h i s i s borne out by the di f f e r e n c e s i n feeding be-haviour sometimes observed i n t h i s study amongst i n d i v i d u a l s of the same species. This p o s s i b i l i t y could best be tested when i t i s possible to grow laboratory cultures of t i n t i n n i d s under completely controlled conditions. 115 T A B L E 24. Species The relationship between tintinnid cell length and number of accumulated food items in two species taken from different experiments. Cell Length Cell Diameter M Lorica Length Number of Food Items Brown Cells 6-8|im dia. Tintinnidium mucicola II II II II 40 40 45 45 80 35 35 35 35 35 40 100 110 125 150 3 7 6 4 11 Volumes of Food Items Um 3  T Y P E 1 T Y P E 2 Natural" large dinos, Isochrysis Eutreptiella sp. galbana Eutintinnus latus 100 70 nil 1450 45 100 70 nil 8500 800 100 70 250 3800 700 100 70 250 7850 840 125 70 220 7700 770 125 70 250 1100 420 200 70 250 6500 525 200 70 275 7000 280 116 (4) Spontaneous changes i n feeding behaviour have not been seen i n t h i s study but Strathmann (1971) has noted them i n echinoderm larvae. (5) The age of a t i n t i n n i d c e l l i s probably not an important cause of v a r i a b i l i t y i n feeding r a t e , apart from the f a c t that a parental c e l l w i l l probably not have fed during d i v i s i o n so that a very newly divided c e l l may contain l e s s food than do others. Table 24 shows that i n two experiments involving d i f f e r e n t species of t i n t i n n i d s there was no r e l a t i o n s h i p between t i n t i n n i d c e l l length and the amount of food contained therein. 117 b) Observations of T i n t i n n i d Motions and Feeding Behaviour Many of the t i n t i n n i d s studied had c h a r a c t e r i s t i c motions so that they could sometimes be i d e n t i f i e d even when without a l o r i c a . A l l t i n t i n n i d s rotate the c e l l and l o r i c a on i t s long axis; and also follow h e l i c a l paths which are normally of a c h a r a c t e r i s t i c angle i n healthy organisms. The largest species are generally the f a s t e s t and several species move at about f i v e c e l l lengths per second. Stensosomella n i v a l i s and Helicostomella k i l i e n s i s move r a p i d l y f o r t h e i r s i z e and Tintinnidium mucicola moves rather slowly f o r i t s s i z e . The most important feature of the motion for a predator of t h i s type i s the frequency of contact with food rather than the v e l o c i t y of the predator. A l l food items tested i n t h i s study were slower i n motion than the t i n t i n n i d s used, and the d i s t r i b u t i o n and motion of both food items and predators can reasonably be regarded as random. There was n e g l i g i b l e con-tact between predators at the concentrations used i n t h i s study. The v e l o c i t y (in um/sec) of a t i n t i n n i d species may be a function of the length and number of i t s adoral c i l i a ; and i t s 'search rate' ( i n ml/hr) i s a function of i t s v e l o c i t y and of the e f f e c t i v e diameter of the vortex created by i t s adoral c i l i a . A h e l i c a l path w i l l not increase the frequency of contact with r a n -domly dispersed prey so long as the predator's v e l o c i t y remains unchanged. Larger and f a s t e r prey are contacted more frequently than smaller and slower prey at the same concentration, and t h i s may explain some of the apparent d i f f e r e n t i a l predation of Tintinnopsis subacuta on E u t r e p t i e l l a sp. when compared to other prey presented s i n g l y i n t h i s study. The 'contact r a t e ' (CR - i n ml/hr) i s the product of the search rate of the t i n t i n n i d species and the concentration ( i n nos/ml) and v e l o c i t y of the prey items. A l l par-t i c l e s which cause a t i n t i n n i d to interrupt i t s normal motion, whether the item i s eaten or not, w i l l reduce the long-term contact rate. Feeding rate 118 (FR) i s calculated e i t h e r : as the number of items observed to be s u c c e s s f u l l y eaten per hour; as the number found to be accumulated i n the t i n t i n n i d per hour: or as the volume of water t h e o r e t i c a l l y cleared of food i n ml/hr., for each type of food. The d i r e c t measurement of t i n t i n n i d v e l o c i t i e s and search rates by eye i s made very d i f f i c u l t by t h e i r h e l i c a l motion and frequent changes of d i r e c -t i o n . Relative and absolute contact rates of t i n t i n n i d s and ( i f they ate) th e i r feeding rates, have been obtained from a study of t h e i r reactions to known concentrations of i d e n t i f i a b l e food items. Unfortunately observations of 'contacts' have a subjective element. Where a t i n t i n n i d ate, or attempted to eat, or changed d i r e c t i o n a f t e r contact - then a 'contact' could be conf firmed. However, a l l t i n t i n n i d species apparently contacted many more par-t i c l e s than were eaten (see Table 26), i . e . p a r t i c l e s would be swept towards the o r a l region and apparently c o l l i d e with the in s i d e of the s p i r a l of adoral c i l i a and be moved out again without being eaten. It i s perhaps i n v a l i d to assume that a t i n t i n n i d was " i n a p o s i t i o n " to eat such items or even that they could have eaten those p a r t i c l e s which cause them to make sudden changes of d i r e c t i o n . Small p a r t i c l e s (4 - 10 jam) were d e f i n i t e l y seen to be eaten at times i n conjunction with a s l i g h t , b r i e f 'tremor' or 'shudder' i n the motioncdf the t i n t i n n i d . The method used by t i n t i n n i d s to r e t a i n small p a r t i c l e s i s unknown. Perhaps mucus may be secreted to 'form a sheet or meshwork near the o r a l region (Laval, 1971); or the p a r t i a l l y overlapping insertionoof the adoral c i l i a may act as a b a r r i e r to any p a r t i c l e which i s intthe centre of the vortex created by the beating of those c i l i a . Strathmann (1971) and Strathmann, e t . a l . , (1972) have shown that many c i l i a r y feeders do not use 119 mucus, but trap p a r t i c l e s either by (a) a l o c a l induced reve r s a l of beat of i n d i v i d u a l c i l i a or (b) by the downstream entrapment of p a r t i c l e s against a second row of c i l i a . T i n t i n n i d s may use one of these or other methods to obtain p a r t i c l e s very small r e l a t i v e to t h e i r o r a l dimensions. Larger items cause a longer i n t e r r u p t i o n of the normal t i n t i n n i d motion; usually with the adoral c i l i a bent inwards so preventing the escape of the prey, whilst the other o r a l c i l i a attempt to push i t further down the cytopharynx. During t h i s period the motionless t i n t i n n i d slowly sinks. This process takes about two seconds f or Tintinnopsis subacuta feeding s u c c e s s f u l l y on E u t r e p t i e l l a sp; and takes 6 - 8 seconds for Eutintinnus l a t u s feeding on the same prey unsuccess-f u l l y . Although p a r t i c l e s smaller than about 4jam. diameter could not be seen with the stereo microscope, one aspect of t i n t i n n i d behaviour indicated that very small p a r t i c l e s weixe be-ihgeeaten /(andwsee Section 4c). A unique feature of t i n t i n n i d s i s the possession of a region of ex t e n s i l e c o r t i c a l cytoplasm adjacent to the o r a l groove/cytopharynx region and c a l l e d the o r a l plug.(see Figure 1). This o r a l plug can be seen o c c a s i o n a l l y to 'pump' or move back and f o r t h quite v i o l e n t l y as the t i n t i n n i d swims i n i i t s normal motion. The exact function of t h i s pumping i s unknown but i t i s p a r t i c u l a r l y obvious i n Tintinnidium mucicola. The pumping motions of f i v e i n d i v i d u a l s of t h i s species were counted as they were observed for varying lengths of time i n seawater containing many f l a g e l l a t e s and other small p a r t i c l e s . 'Contacts' between T_. mucicola and p a r t i c l e s v i s i b l e to the observer were also counted. Both measurements were also made of f i v e i n d i v i d u a l s placed i n seawater which had been f i l t e r e d a few days before but which contained some p a r t i c l e s . The average figures for the t i n t i n n i d s i n the f i l t e r e d water were 7.1 'pumps' and 4.5 contacts/minute; f or those i n the u n f i l t e r e d water the averages were 17.4 'pumps' and 9.7 contacts/minute. Pumps exceeded contacts i n both cases, j u s t 120 as small p a r t i c l e s usually outnumber large ones; and the r a t i o s of pump/con-tact were r e s p e c t i v e l y 1.67 and 1.58 i n the two kinds of water. This would seem to be evidence that the r a t e of 'pumping' at l e a s t i n T_. mucicola i s a d i r e c t function of the number of very small p a r t i c l e s i n the water and may be connected with the r a t e of feeding on them. Unwanted p a r t i c l e s which have d e f i n i t e l y been observed to enter the cytopharyngeal region of a t i n t i n n i d are usually ejected by a momentary r e -v e r s a l of some or a l l of the c i l i a i n the o r a l region. A very powerful or prolonged c i l i a r y r e v e r s a l of t h i s type w i l l cause the t i n t i n n i d to reverse d i r e c t i o n away from the object, as also happens a f t e r contact with objects larger than the t i n t i n n i d . Rarely, a t i n t i n n i d which cannot eject an object i n t h i s way may move i t s aboral end through 180 , and then reverse to move away from the object. Some i n d i v i d u a l c e l l s , p a r t i c u l a r l y of Tintinnidium  mucicola, may show apparent signs of stress i n the form of aberrant motion, for several seconds a f t e r e jecting an unwanted object. The c h a r a c t e r i s t i c s of c e r t a i n large prey items which prevent them from being ingested by a par-t i c u l a r t i n t i n n i d species were experimentally investigated as shown i n Table 25. The predator was Eutintinnus latus ( l o r i c a 250x70 /im; c e l l 80-200x60 /am) and the prey were a l g a l f l a g e l l a t e s from laboratory cu l t u r e s ; E u t r e p t i e l l a  sp. (25x7x4 /am) and Cryptomonas profunda (30x10x4 /im). I t had been noticed i n a q u a l i t a t i v e test that E_. latus contained C_. profunda but not E u t r e p t i e l l a  sp. although the l a t t e r i s only about h a l f the volume of the former. The attempts of .E. latus to ingest these f l a g e l l a t e s when they were pre-sented i n single-prey samples, was observed with the prey i n either normal, or an immobilized condition. Mild sonication for 15 seconds immobilized both prey types, but with d i f f e r e n t r e s u l t s . Sonicated E u t r e p t i e l l a sp. l o s t T A B L E 25. The effect of immobilization by sonication on the successful ingestion of algal flagellates by the tintinnid, Eutintinnus latus. Temperature 1 8 - 2 0 ° C . Rate of* Number Total Successful Handling Contact Number Contact* Ingestion* Prey of Number Ingestion Time with prey prey ce l l s / Rate Rate Prey Type Condition Tintinnids Contacts Events ( S e e s . ) N o . / M i n . mL m l / h r / T i n . m l / h r / T i n . Eutreptiella sp. Normal 3 13 0 6-8 4 6 (Av. 7.3) 5230 0.053 Immobile 2-12 3.0 5230 0.034 0.025 (Av. 4.8) < Cryptomonaa profunda Normal 6 11 6 5-19 2.9 4630 (Av. 9.3) Immobile 3 2? 0 0 0.82 4630? * These rate a are calculated as the total elapsed time of a "run" and Include handling time. 0.038 0.021 0.011? 0? to 122 t h e i r f l a g e l l a e , became more rounded and could not swim, but were d e f i n i t e l y a l i v e and showed the unique euglenoid type of c e l l motion when supported on a glass s l i d e . Sonicated CJ. profunda did not l o s e t h e i r f l a g e l l a e , but were unable to move whether supported or not. (3. profunda may have beeneKilled by t h i s treatment, and the r e s u l t s i n Table 25 may r e f l e c t t h i s . Normal Eutrep- t i e l l a sp. moved at about 250 to 400jam/second at 18-20 C, or about 10 to 16 c e l l lengths/second. j 3 . profunda moved about h as f a s t as E u t r e p t i e l l a sp. 'Contacts with prey' were conservatively equated with v i s i b l e 'attempts at ingestion' since only then was i t c e r t a i n that the t i n t i n n i d s were aware of a prey item. The two contacts with immobilized C^ . profunda were 'glancing blows' causing IS. latus to change d i r e c t i o n s l i g h t l y and may not have been what they seemed, i . e . the r a p i d i d e n t i f i c a t i o n and r e j e c t i o n @f. probably i n -edible prey items. Therefore, i t i s doubtful whether IS. l a t u s rejected the immoblized jT- profunda. However, from Table 25 i t i s obvious that the immo-b i l i z a t i o n of E u t r e p t i e l l a sp. d e f i n i t e l y made a d i f f e r e n c e to the a b i l i t y of IS. l a t u s to handle and eat i t (from 0 to 71% success), and also reduced the handling time by 35%, though t h i s may not be a s i g n i f i c a n t d i f f e r e n c e . Natur-a l l y , random contact with immobilized E u t r e p t i e l l a sp. was l e s s frequent than with normal E u t r e p t i e l l a sp. Normal E u t r e p t i e l l a sp. were always released un-harmed a f t e r attempted ingestion. The very high ingestion rate (or FR/P) of 0.025 ml/hr of IS. l a t u s on immobilized E u t r e p t i e l l a sp. was very s i m i l a r to that on C^. profunda, but the i n d i v i d u a l handling time on the l a t t e r was longer. It i s i n t e r e s t i n g that IS. l a t u s i s about seven-fold larger than T i n t i n - nopsis subacuta (see Table 1) which ingests E u t r e p t i e l l a sp. i n about two seconds. Both these t i n t i n n i d s and Tintinnopsis c y l i n d r i c a have adoral c i l i a of about 35 jam length, but T_. c y l i n d r i c a moves a l i t t l e more slowly than 123 T_. subacuta and was not seen to ingest E u t r e p t i e l l a sp. i n the laboratory. Stenosomella ventricosa has exceptionally thick and powerful-looking adoral c i l i a (Table 1) but moves only a l i t t l e f a s t e r than T_. subacuta and eats E u t r e p t i e l l a sp. much l e s s frequently than the l a t t e r (see Section 4a ( i i ) ) . However, the adoral c i l i a of S^  ventricosa are 50jam long; as are those of F a v e l l a serrata which i s about 10 times l a r g e r . Perhaps the number of c i l i a per t i n t i n n i d rather than t h e i r length i s important i n providing both higher speeds and increased a b i l i t y to subdue prey. I t was impossible to make de-f i n i t i v e counts of the numbers of adoral c i l i a i n t h i s study, but they ap-peared to d i f f e r rather narrowly between species from about 16 to 24. T i n t i n n i d contact rates Table 26 shows the contact rates of various t i n t i n n i d species on natural and laboratory food items i n the same, and i n d i f f e r e n t , experiments. Other contact rates may be seen i n Table .8 and 9 (accumulation experiments). These r e s u l t s i n d i c a t e that under the conditions of observation and with the c e l l concentrations used, t i n t i n n i d s ate very few of the p a r t i c l e s encountered even when the t i n t i n n i d s were ac t i v e and i n large numbers i n the environment. Therefore i t i s possible that the experimental conditions were not optimal fo r t i n t i n n i d feeding. I t can also be seen i n Table 26 that the a c t i v i t y of d i f f e r e n t species i s not a f f e c t e d to the same extent by dense concentrations of p a r t i c l e s . Although there i s considerable v a r i a b i l i t y between experiments, on the whole the values of the contact rates (and therefore probably the search rates) for Tintinnopsis subacuta and Stenosomella ventricosa are s i m i -l a r , with the l a t t e r perhaps a l i t t l e higher. This supports the r e l a t i v e feeding rates estimated i n the r e s u l t s of the Accumulation (Section 4a) and Coulter Counter (Section 4c) experiments, except when the two species were T A B L E 26. Observed contact rates of various tintinnid species on natural and laboratory food it ems. Experiment Number P r e / , Predator Average contacts/ Prey m i n / T i n . N o s / m l Average CR Ingestions/ F R Relative m l / h r / T i n . m i n / T i n . m l / h r / T i n . CR Comments 1A Natural T . subacuta 18.8 12; 200 0.092 ni l nil IF 1.0 particles II T . parvula 21,8 12, 200 0.107 n i l nil 1.2 II T . mucicola 3.5 12, 200 0.017 ni l ni l 0.2 II S. nivalis 28.8 12, 200 0.141 ni l ni l 1.5 IB Natural T . subacuta 22.4* 19, 700 0.068 ni l ni l IF 1.0 particles and S. ventricosa 28.6 19, 700 0.087 ni l n i l ' 1.3 Dunaliella T . mucicola 15.8 19, 700 0.048 0.75 0.0023 0.7 tertiolecta II S. nivalis 18.5 19. 700 0.056 n i l ni l 0.8 * 0.29 of contacts were with D. tertiolecta (0.37 of a l l particles) IB Natural T . subacuta 16.3 ? nil ni l . I F 1.0 12 hrs) particles T . parvula 11.3 - - ni l nil 0.7 and S. ventricosa 17.7 - - ni l n i l 1.1 Dunaliel la T . mucicola 7.0 - - ni l n i l 0.4 tertiolecta S. nivalis 6.3 - - nil ni l 0.4 2 Natural T . subacuta 36.2 9. 500 0.226 ni l ni l I F 1.0 particles S. ventricosa 40.0 9. 500 0.252 ni l ni l 1.1 moat < 10 T . mucicola 22.0 9. 500 ? nil nil 0.6 u m S. nivalis 22.0 9. 500 ? ni l ni l 0.6 3 Natural T . cyl indrica 32.0 71,000 0.027 ni l ni l IF 1.0 particles T . mucicola 18.2 71,000 0.022 2.30 0.0018 0.8 H . kiliensis 42.2 71,000 0.036 ni l ni l 1.3 E. tubulosus 9.8 71,000 0.011 0.37 0.0004 0.4 4A Natural T . cyl indrica 27.0 n i l I F 1.0 particles T . mucicola 9.0 - _ 0.8 0.3 and H . kiliensis 7.9 - - nil 0.3 dense E. latus 14.2 - - ni l 0.5 Unusually sluggish Rejections lengthy Very active Very slow, shallow spirals Violent rejection motion Rapid shallow spirals Bloom ~-6/ml Isoselmis sp. T A B L E 26. Continued Experiment N i l m b o r Prey Predator Average contacts/ ' Prey min/Tin . Nos /ml CR m l / h r / T i n . Average Ingestions/ m i n / T i n . . F R m l / h r / T i n . Relative CR Commonta 4 B Natural particles and dense Dunaliella tertiolecta T . cylindrica T . mucicola H . kiliensis E . tubulosus 29.0 9.2 8.2 2.0 nil n i l ni l n i l IF 1.0 0.3 0.3 0.1 Monochrysis lutheri T . parvula T . mucicola 18.4 13.9 36,000 36, 000 0.030 0.023 ni l 0.80 0.0017 IF 1.0 0.8 126 eating E u t r e p t i e l l a sp.. The contact rates of Tintinnopsis parvula, T i n t i n - nidium mucicola and Stenosomella n i v a l i s were on the whole not very d i f f e r e n t , and were about 0.3 to 1.0 of the values f o r T_. subacuta. This also p a r t l y supports the r e l a t i v e feeding rates calculated by the other methods, p a r t i c -u l a r l y f o r T_. parvula. Tintinnopsis c y l i n d r i c a appeared to have a s l i g h t l y lower contact rate than T_. subacuta when seen i n the same sample ( q u a l i t a t i v e observation)'-but there are no data f o r t h i s comparison. T_. mucicola and HelicostdmeTla  k i l i e n s i s had contact rates which were s i m i l a r and equivalent to 0.3 to 1.3 of those of T_. c y l i n d r i c a . The contact rates of Eutintinnus tubulosus were only about 0.1 to 0.4 of those of T_. c y l i n d r i c a and l e s s than those of H. k i l i e n s i s (and see Table 9). In contrast, the numbers of accumulated food items iiri" E_. tubulosus and H. k i l i e n s i s were f a i r l y s i m i l a r i n one accumu-l a t i o n experiments (see Table 9), but there are no short-term accumulation estimations of feeding rate i n these two species. The large t i n t i n n i d E u t i n - tinnus latus i n one experiment i n Table 26 has a contact rate of only about h a l f of that of T_. c y l i n d r i c a , although i n the r e s u l t s seen i n Table 25 E. l a t u s had a very high feeding rate. Table 26 shows that very few items were d e f i n i t e l y seen to be eaten during these observations, but the feeding rates shown for T_. mucicola are comparable with those seen i n the r e s u l t s of the accumulation experiments (section 4a') . Attempts were made to observe changes i n contact and feeding rates during the progress of experiments and as t i n t i n n i d s starved or slowed t h e i r apparent accumulation of food items, but none of these attempts was s a t i s f a c t o r y . 127 c) The e f f e c t of microzooplankton on natural and laboratory  phytoplankton populations (Coulter Counter Experiments) General The feeding rates of two species of t i n t i n n i d s and the l a t e naupliar (Stage V and VI) stages of a small copepod (probably Pseudocalanus minutus) and a barnacle are shown i n Appendices 1 to 11. The regulatory e f f e c t of the feeding of Tintinnopsis subacuta on natural food p a r t i c l e concentrations i s shown i n Fi g s . 12, 13 and 14. The complete l i s t s of observations; multiple c o r r e l a t i o n c o e f f i c i e n t s f o r 10 v a r i a b l e s ; and simple l i n e a r regression co-e f f i c i e n t s on v a r i a b l e (1) and plots f o r these Coulter Counter experiments as c a l c u l a t e d with the Uni v e r s i t y of B r i t i s h Columbia Computer programme STRIP may be found i n Appendices 1 to 11. A l l r e s u l t s were adjusted to an experi-mental duration of 24 hours before a n a l y s i s . Tintinnopsis subacuta was used i n experiments with n a t u r a l l y occurring p a r t i c u l a t e material (44 obser-vations) and laboratory phytoplankton cultures (15 observations); and Stene somella ventricosa was used with laboratory cultures only (16 observations). Crustacean n a u p l i i were used mainly with laboratory cultures (5 observations), but one experiment was c a r r i e d out with a f i e l d sample. There i s one Coulter Counter r e s u l t of feeding by Tintinnopsis parvula and another by the holotrich. c i l i a t e T i a r i n a fusus, both on f i e l d samples. The l a t t e r three s i n g l e ex-periments were included i n the analyses of c o r r e l a t i o n and regression only i n the ' a l l samples' category. One experiment involving two species of r o t i f e r s has been analysed separately (Table 28). Some of the observations included i n the analyses were ' r e p l i c a t e s ' i n the same experiment. Where two or three p a i r s of con t r o l and experimental f l a s k s were used, every possible value of feeding rate (FR/P) 128 was calculated from every possible combination of a l l values of C^, , and E2 and included i n the analyses to increase the t o t a l number of obser-vations. S i m i l a r l y , f l a s k s within a p a r t i c u l a r experiment which were subject to manipulation such as placement i n the l i g h t or dark, a d d i t i o n a l predators, etc., have a l l been included without d i s t i n c t i o n i n the general analyses. Calculations of feeding rate as net t o t a l consumption (NTC) or as edible spectrum only (ESO) are shown separately. It i s obvious from the l i s t of observations i n the Appendices that there i s great v a r i a b i l i t y i n the food consumption rate ER'/P (or v a r i a b l e (1)) even amongst ' r e p l i c a t e s ' i n the same experiment (see General Discussion). Mu l t i p l e c o r r e l a t i o n c o e f f i c i e n t s . Only those c o r r e l a t i o n c o e f f i c i e n t s greater than 0.5 have been included i n Tables 27 a,b,c,d. The remainder of the c o r r e l a t i o n matrices may be seen i n Appendices 1 to 11. Those pairs of v a r i a b l e s which had c o r r e l a t i o n coef-f i c i e n t s greater than 0.5 are not n e c e s s a r i l y the same either i n the NTC and ESO r e s u l t s f o r any one subset of observations (e.g. Tintinnopsis subacuta on laboratory food); nor for NTC (or ESO) r e s u l t s as compared between subsets of observations. Also, the sign of the c o r r e l a t i o n i s often d i f f e r e n t f o r the same pair of va r i a b l e s i n NTC and ESO r e s u l t s , and/or between subsets of observations. C o e f f i c i e n t s f o r ESO r e s u l t s are generally higher than those f o r NTC r e s u l t s . Very few v a r i a b l e s are strongly correlated with the food consumption rate (variable 1), and by far the best f i g u r e s are found with E^ (variable 2) and logmean E (3) i n Tintinnopsis subacuta feeding on laboratory cultures of phytoplankton (Table 27b). The greater coherence of these data may be l a r g e l y explained by the fac t that i n most cases the same phytoplankton 129 T A B L E 27A. Multiple correlation coefficients from Coulter Counter experiments. (only those > 0.5 are shown) Tintinnopsis subacuta on natural samples. Variable Net total consumption (NTC)  Edible spectrum only (ESO)  (1) F R / P Feeding rate l im 3 /pred/hr 39 observations N i l 44 observations 2 (.59), 3(.62) (2) E i Initial experimental particle volume lirrrV ml 3(.91), 9(-.8l) 1(.59), 3(.98), 4(.68) (3) Log mean E 12 hr. log mean experimental particle volume Uim3/ ml 2 ( . 9 D , 8(.71), 9(-.63) 1(.62), 4(.75) (4) ESP. Number of size classes with consumption 5(.58) 2(.68), 3(.75) (5) ^ / Q Increase in total control volume in 24 hrs. 4(.58) N i l 130 T A B L E 27B. Multiple correlation coefficients from Coulter Counter experiments. (Only those > 0.5 are shown) Tintinnopsis subacuta on laboratory food. Variable Net total consumption Edible spectrum only (NTC) (ESO)  (1) F R / P Feeding rate l im 3 /pred/hr (2) E i - Initial experimental volume | i m 3 / m l (3) Log mean E 12 hr log mean experimental particle volume Jim 3 / ml (4) ESP - Number of size classes with consumption 10 observations 6(-.67). 9(-.94) 3(.99). 7(.73), 8(.83) 2(.99). 7(.67), 8(.81) 15 observations 2(.96). 3(.92), 7(.50) 1(.96), 3(.97), 4(.52), 7(.51) 1(.92), 2(.97) Nil (5) Ca /C x - Increase in 7(-.6l) .total control volume (6) PCON - Number preds/ ml (7) Alive - Estimated live fraction of preds at 12 hrs (8) Temperature °C (9) Sal. - Salinity & (10) Time - Total duration of experiment l(-.67), 9(.59) 2(.52) 7(-.59) 7(-.55) 2(.73), 3(.67), 5(-.6l) 1(.50), 2(.51), 5(-.59) 8(.80) 6(-.55), 9(-.51) 2(.83), 3(.8l), 7(.80) l(-.94), 6(.59) Nil Nil 7(-.51), 10(-.51) 9(-.51) T A B L E 27C(i). Multiple correlation coefficients from Coulter Counter experiments. Only those > 0.5 are shown. Stenosomella ventricosa (all values) on laboratory food. T A B L E 27C(ii). Stenosomella ventricosa (one high value of F R / P omitted) on laboratory food. Variable Net total consumption (NTC) 1 3 observations Edible spectrum only (ESO) 16 observations (1) F R / P - Feeding rate - um^/pred/hr (2) E i - Initial experimental volume um^/ml (3) Log mean E - 12 hr mean exp. vol. (jtm^/ml (4) E S P - Number of size classes with consumption (5) C a / C i - Increase in total control volume (6) P C O N - Number predators/ml (7) Al ive - Estimated live fraction of predators at 1 2 hrs (8) Temperature ° C (9) Sal. - Salinity*, (10). Time - Total duration of experiment 6(-.67) 3(.60), 5(-.62), 7(-.50) 8(.8l) • 6(-.57), 7(-.67). 10(-.51) 3(.90). 4(.71) 2(.60). 9(-.58), 10(.52) 2(.90). 4(.55). 10(.©4) Nil 2(.71). 3(.55) 2(-.62). 7(.79), 8(-.90) 7(.64). 8(-.74) 10(.54) K-.67), 9(.67) l(-.57). 9(.62) 2(-.50). 5(.79). 8(-.81) l(-.67). 5(.64). 8(-.77) 10(.68) 10(.69) 2(.8l). 5(-.90). 7(-.81) 5(-.74). 7(-.77) 3(-.58), 6(.67), 10(r.53) 6{.62), 10(-.54) 3(.52), 5(.54), 7(.68). , 1(-.51), 3(.64), 7(.69) 9(-.53) ' 9(-.54) Edible spectrum only (ESO) 1 5 observations 2(.56), 3(.56). 6(-.6l) 1(.56), 3(.92). 4(.71) 1(.56), 2(.92). 4(.54). 10(.58) 2(.71). 3(.54) 7(.75), 8(-.72) '(-.61). 9(i68), 10(-.57) 5(.75). 8(-.94) 5(-.72). 7(-.94) 6(.68). 10(-.70) 3(.58), 6(-.57). 9(-.70) T A B L E 27D. Multiple correlation coefficients from Coulter Counter experiments. (Only those > 0.5 shown) Barnacles and copepod Nauplii on laboratory food. Variable Net total consumption (NTC) Edible spectrum only (ESO) (1) FR/P - Feeding rate - u m 3 / p r e d / h r (2) E x - Initial experimental volume u m ^ / m l ^3) Log mean £ - 12 hr log mean exp. vol. Mm 3 /ml (4) E S P - Number of oize classes with consumption (5) C a / C i - Increase i n total control volume (6) P C O N - Number predator s / m l 3 observations 5 observations 2(-.73). 3(-.90). 4(.84) 2(.82), 3(.75). 4(.78), 8(-.64) 5(-.97), 6(-.6S). 7(.98) 9(-.68) 8(-.91). 9h94) K-.73) , 3 (.96). 4(-.98). 5(.87). 7(.85) 8(.95). 10(.91) K-.90) , 2(.96). 4(-.99) 5(.98). 7(.97), 8(1.0) 9(,68). 10(.75) 1(.82). 3(.99). 4(.79) 1(.75). 2(.99). 4(.70) K-.97) . 2(.87). 3(.98) 1(.78), 2(.79). 3(.70). 8(-.73) 5(-.95), 7(-.93). 8(-.99) 10(-.72) 9(-.60), 10(-.82) 1 (-.97. 2{.87). 3(.98). 4(-.94). 7(1.0), 8(.98) 9(.83). 10(.58) 6(.80). 7(.92), 9(.80) l(-.68), 7(.52), 9(.90) 5(.80). 7(.66), 9(.90) (7) Alive - Estimated live fraction of predators at 12 hrs (8) Temperature ° C (9) Sal. - Salinity & (10) Time - Total duration of experiment l(-.98). 2(.85), 3(.97) 4(-.93), 5(1.0). 6(.52) 8(.98), 9(.85), 10(.56) K-.91). 2(.95), 3(1.0) 4(-.99), 5(.83). 6(.90) 7(.85). 9(.71) K-.94). 3(.68). 4(-.60) 5(.83). 6(.90),. 7(.85) 8(.71) 2(.9D. 3(.75). 4(-.82) 5(.58). 7(.56) 5(.92). 6(.66), 9(.6l) l{-.64). 4(-.73). 10(.77) l(-.68). 5(.80). 6(.90). 7(.61) 4(-.72), 8(.77) 133 species was used, namely Monochrysis l u t h e r i either s i n g l y , or with one other species. The strong c o r r e l a t i o n s seen i n the r e s u l t s of experiments with the crustacean n a u p l i i are n o n - s i g n i f i c a n t (Table 27d) due to the small number of observations. Food consumption rates (variable 1) were usually most strongly correlated with values (variable 2) and Logmean E values (3) as expected. More s u r p r i s i n g l y , c o r r e l a t i o n c o e f f i c i e n t s greater than 0.5 were found between i n d i v i d u a l feeding rate (variable 1) and predator concentration (6); f r a c t i o n of predators a l i v e a f t e r 12 hours (7); and s a l i n i t y (9). There i s no obvious explanation for these l a t t e r r e l a t i o n s h i p s . On the other hand, the expected strong c o r r e l a t i o n between feeding r a t e (variable 1) and the number of s i z e classes showing net consumption or 'ESP' (variable 4) and between (1) and the growth rate of c o n t r o l populations (V^/C^ ~ v a r i a b l e 5) di d not appear (but see Materials and Methods). However, v a r i a b l e (4) was i n some cases strongly and p o s i t i v e l y correlated with E^ values (variable 2). This implies that predators i n r e l a t i v e l y high concentrations of food d i d not r e s t r i c t t h e i r consumption to a narrower portion of the food s i z e spectrum than d i d those i n r e l a t i v e l y low concentrations of food (and see E l e c t i v i t y r e s u l t s i n t h i s Section). That i s , — i f one ignores the unknown p r i o r nut-r i t i o n a l h i s t o r y of the p r e d a t o r — they do not become 'choosier' of the s i z e of t h e i r food when s a t i a t i o n i s becoming more l i k e l y or vice-versa, — a t l e a s t at these l e v e l s of concentration of food, which a l l seem to be below the OFC (see below). There was s u r p r i s i n g l y , no consistent r e l a t i o n s h i p between tem-perature (variable 8) , and food consumption rate (1) , 'width of- spectrum' eaten (4), or C^/C^ (5). Linear regression c o e f f i c i e n t s Simple l i n e a r regressions were calculated f or a l l other variables on 2 feeding rate (variable 1). Regression c o e f f i c i e n t s and r values f o r a l l 134 v a r i a b l e s and computer plots for v a r i a b l e s 2,3 and 4 are shown i n Appendices 1 to 11. 2 As i n the case of the c o r r e l a t i o n c o e f f i c i e n t s , the r values for the ESO r e s u l t s were generally higher than those for the NTC r e s u l t s . The values of the regression c o e f f i c i e n t s of v a r i a b l e s (2) and (3) on the food consumpr1 -t i o n r a t e (1) were generally greater, and the regression l i n e was more s i m i l a r to the form expected, f o r the ESO r e s u l t s than f o r the NTC r e s u l t s . This seems to confirm that the ESO method of c a l c u l a t i n g food consump-t i o n rates may be more useful for microzooplankton than the more l o g i c a l NTC method. The (ESO) calculated regression l i n e s of values of (variable 2) or Logmean E (variable 3) on feeding rate (variable 1) showed the following features: (i ) a negative intercept on (1) — i . e . when v a r i a b l e (1) = 0, then v a r i a b l e (3)>0. ( i i ) a p o s i t i v e slope; and ( i i i ) no asymptote apparent i n the data. (i ) negative intercept - or threshold feeding-*value 'Threshold' feeding values, or the lower l e v e l of t o t a l a v a i l a b l e food at which feeding (apparently) stops, are a common feature with some planktonic crustacean f i l t e r feeders (Adams and Steele, 1966J); Parsons e t . a l . , 1967) .and with some planktivorous f i s h (Parsons and Lebrasseur, 1970). Parsons e t . a l . (1967) and Poulet (1974) also u t i l i z e d the Coulter Counter technique. Feeding thresholds applicable over the t o t a l s i z e range of the biomass of food a v a i l -able with one feeding technique are probably useful (or e s s e n t i a l ) i n enabling the predator under natural conditions to avoid wasteful e f f o r t when the energy return i n feeding with that technique i s n e g l i g i b l e . Poulet (1974) found that 135 the small copepod Pseudocalanus minutus showed a feeding threshold for each s i z e category and could switch i t s feeding emphasis away from a s i z e c l a s s containing a low t o t a l volume of p a r t i c l e s . Despite the i n s u f f i c i e n t d e t a i l of experimental technique, and the apparently inadequate methods of c a l c u l a t i o n as given by Poulet (1973 and 1974), any feeding behaviour of t h i s type would be a very useful a t t r i b u t e for an organism r e l i a n t on a food supply highly v a r i a b l e i n i t s s i z e composition; and would a i d the maintenance of e c o l o g i c a l s t a b i l i t y ( i f any) i n the plankton. The mechanisms(s) used by crustacea to detect these differences i n t o t a l volume between siz e classes i s d i f f i c u l t to imagine. The Coulter Counter r e s u l t s i n t h i s study show low apparent feeding thresholds f o r microzooplankton which may be experimental a r t i f a c t s and'which are (as always) l i n e a r extrapolations from known values. They are p a r t i c u l a r l y suspicious i n t i n t i n n i d s as i t would seem impossible for t i n t i n n i d s to stop feeding and yet continue swimming. It i s i n t e r e s t i n g that the larvae of several species of f i s h which eat food items i n d i v i d u a l l y do not show feeding thresholds. Parsons et^.al^. (1967), Poulet (1974) and others have shown that planktonic crustacea also have a q u a l i t a t i v e lower s i z e feeding threshold. The a t t r i b u t e s of t i n t i n n i d s i n t h i s respect are dealt with i n Section 4a. Lower threshold feeding values calculated from the p l o t s i n Appendices 1 to 11, and the corresponding regression c o e f f i c i e n t s , are shown i n Table 28. Over a l l experiments, the extrapolated threshold values of Logmean E range 3 3 from 35 to 340 x 10 um /ml, with no clearcut differences i n the range of values between the t i n t i n n i d species or between t i n t i n n i d s and n a u p l i i (Table 3 3 28). The threshold value for ' a l l samples' (ESO) i s 72 x 10 um /ml — equi-valent i n volume to approximately 1440 Monochrysis l u t h e r i c e l l s , or 144 E u t r e p t i e l l a sp. c e l l s / m l . These values are very low indeed, e s p e c i a l l y as 136 T A B L E 28. Microzooplankton lower threshold feeding values and regression coefficients of Log mean E (variable 3) when food consumption rate (variable 1) is zero . . N.S. = non significant at .0 5 probability level. Species of predator Eood type Calculation method Lower Log mean E (3) Thresholds (Um 3 x 10 3 /ml) Regression Coefficient of Variable (3) on Variable (1) Tintinnopsis subacuta II Natural Natural N T C E S O 35 124 0.0197 0.0377 II II Laboratory Laboratory N T C ESO Intercept Negative 101 0.0033 (N.S.) 0.0205 Stenosomella ventricosa II II Laboratory Laboratory Laboratory N T C E S O ESO 340 Regression Negative 46 (one high value omitted) 0.0068 (N.S.) Negative 0.0089 Barnacle and copepod/ Nauplii II Natural Laboratory Laboratory E S O N T C ESO Regression Negative 38 0.0093 (N.S.) Negative • 0.0238 (N.S.) Tintinnopsis parvula Natural E S O -- 0.0056 (N.S.) T iar ina fusus Natural ESO 0.231 (N.S.) A l l samples N T C E S O Intercept Negative 72 0.0015 0.0247 137 they represent a l l p a r t i c l e s l e s s than approximately 20yim diameter. 3 6 Using the carbon pug) to volume tym x 10 ) conversion f a c t o r (.052) 3 3 given by Parsons e_t.al_. (1967), a threshold value of 72 x 10 /im /ml i s equivalent <tb 0.0037ug/carbon/ml. The l a t t e r i s much lower than the thres-hold values Poulet (1973) gives f o r Pseudocalanus minutus i n a coa s t a l em-bayment and only l/14th the value of the lowest of those given f o r three species of coa s t a l planktonic crustaceans by Parsons et.al^. (1967). I t i s almost c e r t a i n l y only c o i n c i d e n t a l that the average threshold value i s about equivalent to the carbon content of one Tintinnopsis subacuta (see Section 2). ( i i ) p o s i t i v e slope of regression Although the regression analysis used presupposes a s i n g l e l i n e a r r e -l a t i o n s h i p between v a r i a b l e s (1) and (3), there are few data points which ob-vi o u s l y v i o l a t e t h i s assumption i n the various sub-sets of observations. An exception to t h i s i s seen i n the r e s u l t s of Tintinnopsis subacuta feeding on natural food, where some of the values of an experiment involving a natural 'bloom' of E u t r e p t i e l l a sp. appear to be unusually high and l i e perhaps on a d i f f e r e n t , or on an exponential curve (Appendix 1 ) . However, other ' r e p l i -cate' r e s u l t s were much lower, and there was considerable 'scatter' amongst the r e s u l t s i n that experiment. Several of the NTC values show no p o s i t i v e l i n e a r i t y , which lends sup-port to the s u r p r i s i n g u t i l i t y of the ESO method of c a l c u l a t i o n , but which makes d i f f i c u l t any approximate p r e d i c t i o n of the feeding r a t e of a species from a knowledge of merely the i n i t i a l t o t a l p a r t i c l e volume i n a sample. The values of the regression c o e f f i c i e n t s are equivalent to an u n r e a l i s t i c a l l y continuous and uniform i n d i v i d u a l feeding r a t e of between 0.33 and 3.8% of the 138 p a r t i c l e volume/ml/hour (and see Section 4a and 4b); with an exceptional s i n g l e value of 23.1% for the h o l o t r i c h c i l i a t e T i a r i n a fusus (ESO only). Tintinnopsis parvula (1 experiment only) shows a small c o e f f i c i e n t (0.56%/ ml/hr) and t h i s t i n t i n n i d has a c e l l volume and a 'search rate' both about h a l f of that of T_. subacuta ( c o e f f i c i e n t of 0.33 to 3.8%/ml/hr) and Stenosomella ventricosa ( c o e f f i c i e n t of 0.68 to 0.89%/ml/hr). The l a t t e r two species are s i m i l a r to each other i n both c e l l s i z e and 'search r a t e ' (see Sections 4a and 4b) . However some feeding rates of T_. parvula on small prey items shown i n Section 4a are as high as or higher than those of T_. subacuta. The crustacean n a u p l i i show a s i m i l a r range of values at 0.93 to 2.38%/ml/hour although larger than t i n t i n n i d s . The c o e f f i c i e n t s calculated for t i n t i n n i d s i n the short-term accumulation experiments (Section 4a ( i i ) ) are a l l 1.1%/ml/hr or l e s s . A l l these values are about 10-fold lower than thenmaximum i n d i v i d u a l feeding c o e f f i c i e n t s given by Poulet (1973) for the much larger adult copepod Pseudocalanus minutus (17%/ml/hr) over a food s i z e range from 2 to 114 um. Feeding c o e f f i c i e n t s of two species of planktonic r o t i f e r s (see the end of t h i s Section) ranged from 0.29% (NTC) to 5.53%/ml/hr (ESO) (and see General Discussion). ( i i i ) lack of asymptote I t i s obvious from the computer plots'of the regression of Logmean E on feeding rate (FR/P) i n Appendices 1 to 11 that there i s no case of a trend i n the data towards a p o s i t i v e c u r v i l i n e a r r e l a t i o n s h i p or towards an asymp-tote. Since t h e o r e t i c a l l y there must be an asymptote, or at l e a s t a curvature for each predator at some high concentration of a v a i l a b l e food, i t must be concluded that these (ESO) Coulter r e s u l t s occur below such optimum l e v e l s of e d i b l e p a r t i c u l a t e material (OFC) (but see Section 4a ( i i ) ) . The d i s c r e -pancy between the l e v e l s of the OFC shown by the two experimental methods 139 i n t h i s study i s in e x p l i c a b l e . The maximum p a r t i c l e concentrations used (between 1 and 20 pm diameter) are equivalent to 1.92 ppm (by volume) f or laboratory cultures and 0.76 ppm for natural samples (see also General Dis-cussion). However, the feeding rate asymptote for Pseudocalanus minutus according to Parsons et_.al. (1967) occurs at about 11.0 ppm of p a r t i c l e s between 1 and 114 um diameter. The Feeding E l e c t i v i t y of Ti n t i n n i d s i n Coulter Counter Experiments Indices of e l e c t i v i t y or feeding preference are i n most cases merely indices of the r e l a t i v e ease with which the food items i n question are caught by a predator at any time. One measure which does not confuse c a t c h a b i l i t y with preference or s e l e c t i o n i s the preference c o e f f i c i e n t of Rapport e t . a l . (1972). In order to c a l c u l a t e t h i s c o e f f i c i e n t one must know the feeding rate of the predator on the food i n question i n sing l e and i n mixed cultures, at equivalent d e n s i t i e s . Rapport et^._al_. (1972) and Rapport (unpublished) have calculated preference c o e f f i c i e n t s f o r the c i l i a t e Stentor coeruleus on several foods. For the purpose of estimating the possible d i f f e r e n t i a l feeding rates of t i n t i n n i d s on^various s i z e classes of p a r t i c l e s i n these Coulter Counter experiments, Ivley's e l e c t i v i t y index (here c a l l e d E l ) has been used. Ivlev's index was chosen since there were no comparative cases of predation on si n g l e and mixed food cultures i n these experiments. E l = r ± " P l (5) r i + p i where r i = the proportion of item i i n the d i e t of the predator and p i = the proportion of item i i n the environment. Therefore, i n t h i s section E l i s a measure simply of the a v a i l a b i l i t y and ease of capture of an item or 140 Figure 8. Relationship between e l e c t i v i t y values (El) of Tintinnopsis subacuta on natural p a r t i c l e s and the mean diameter of Coulter Counter s i z e c l a s s e s . (1)(2)(3)(4) (5) (6) (7) (8) (9) ( 1 0 ) P A R T I C L E D I A M E T E R (um) 142 s i z e c l a s s of p a r t i c l e s and not a measure of i t s s e l e c t i o n or preference by the t i n t i n n i d . E l has the t h e o r e t i c a l l i m i t s of -1.0 to +1.0. A zero r e s u l t indicates that a s i z e c l a s s of p a r t i c l e s i s eaten exactly i n the pro-portion i n which i t occurs i n the environment. A p o s i t i v e E l f r a c t i o n i n d i -3 cates that the feeding rate (inyum / h r / t i n t i n n i d ) on a p a r t i c u l a r s i z e c l a s s i s a larger proportion of the t o t a l net feeding r a t e i n that experimental r e p l i c a t e , than the logmean E value of that s i z e c l a s s i s of the t o t a l logmean E. value (also see Section 4a). Figures 8 to 11 show the r e l a t i o n s h i p of the e l e c t i v i t y i n d i c e s with other f a c t o r s for each of 10 s i z e c l a s s e s from that of mean diameter 1.78 jam (class 1) to that of mean diameter 14.3 yum (class 10). Only the values of Tintinnopsis subacuta feeding on .natural samples have been used. Figure 8 shows the r e l a t i o n s h i p between E l and s i z e c l a s s ; Figure 9 shows the r e l a t i o n s h i p between E l and the mean t o t a l p a r t i c l e volume (logmean E) for each s i z e c l a s s ; Figure 10 shows the r e l a t i o n s h i p be-tween E l and the t o t a l logmean E f o r a l l s i z e classes i n the experiment; and Figure 11 shows the r e l a t i o n s h i p between the E l values and the index of increase of p a r t i c l e volume i n the c o n t r o l vessel (G^/C^) for each s i z e c l a s s . In Figures 9,10 and 11 the logmean E and O^/C^ values have been r e c a l c u l a t e d for an experimental duration of 24 hours. There i s no simple r e l a t i o n s h i p between p a r t i c l e s i z e and e l e c t i v i t y index (Figure 8). P o s i t i v e and negative values are found i n almost a l l s i z e c l a s s e s , except 6 (5.66 yum), but the values are c o n s i s t e n t l y p o s i t i v e only in those s i z e classes from 3.57 to 7.12 yum mean diameter. Size classes both larger and smaller than t h i s have a wider range of E l values, both p o s i -t i v e and negative. Size classes 1,2 and 3 (1.78 to 2,82 pm) are more often negative than p o s i t i v e and the two values i n c l a s s 10 (14,3 u^m) are both p o s i -t i v e . Therefore, the o v e r a l l trend i s for e l e c t i v i t y values for Tintinnopsis 143 Figure 9. Relationship between e l e c t i v i t y values (El) of Tintinnopsis subacuta on natural p a r t i c l e s and the Logmean E values of each Coulter Counter s i z e c l a s s . 144 o -oo 00 1 0 —O-O 00 l"vt-« oo t^po o •t-o o CN <N CS CM " I N n CN CN 0- CO N O- C O , 145 Figure 10. Relationship between the e l e c t i v i t y values (El) of Tintinnopsis  subacuta on natural p a r t i c l e s and the t o t a l Logmean E values of a l l Coulter Counter s i z e classes. 10 10 1 1 8 7*8 -3_ 1 7 4 4 1 J 6 •8 9 6 67 4 fl? 3 46 ? & 4 j J 4 7* 87 7 9 7 98 8 8 9 7 5 8 3 2 3 9 3 2 • 3 23 3 S 3 2 2 2 8 2 2 8 6 5 8 9 8 ? 76 4 8 4 4 53 2 4 94 4 . 5 2 4 82 240 260 280 300 320 340 360 380 400 420 440 460 480 500 520 540 560 580 600 620 640 660 680 700 TOTAL LOGMEAN E (pm3 x 103) 147 subacuta to be more c o n s i s t e n t l y p o s i t i v e i n Coulter s i z e classes larger than about 3 pm, but not to increase i n magnitude with s i z e above t h i s . Par-t i c l e s as small as 1.6 u^m are eaten to some extent by t h i s species. In Figure 9 i t can be seen that although the mean t o t a l volume of p a r t i c l e s i s greatest i n the smaller s i z e c l a s s e s , there i s no tendency for e l e c t i v i t y values to increase with the t o t a l p a r t i c l e volume i n each s i z e c l a s s . Figure 11 shows that the p o t e n t i a l rate of increase i n the number of p a r t i c l e s per s i z e c l a s s (C2/C^); may be r e l a t e d i n a complex manner to the e l e c t i v i t y index of a s i z e c l a s s . There i s no r e l a t i o n s h i p between C^/C^ a n c * ^ w n e n ^2^1 i s equal to or l e s s than 1.0, i . e . when there was no change, or a decline i n the t o t a l p a r t i c l e volume f o r that s i z e c l a s s i n the control v e s s e l . In the majority of cases, t o t a l C^/C^ values greater than 1.0 ( i . e . showing 'growth') are associated with s i z e classes with p o s i t i v e E l values. The greatest 2^^ C"1 values occur i n s i z e classes 5 and 6 (between about 4 and 6.5 ^im) , but these s i z e classes do not have the l a r g e s t p o s i t i v e E l values (see also Figure 8). There i s not complete independence between these factors since the E l values are based upon feeding r a t e s ; and the c a l c u l a t i o n of the l a t t e r i n -volves the C ^ ^ i r a t i ° > a n a t n e logmean E value for each s i z e c l a s s (see Section 3b ( i v ) ) . In general, the sign of the e l e c t i v i t y index i s consistent only i n the middle range of the s i z e classes i n these Coulter Counter experi-ments, and these s i z e classes are those which show not the greatest mean part-i c l e volume, but the greatest growth rate i n c o n t r o l s . The magnitude of the e l e c t i v i t y index of Tintinnopsis subacuta bears no consistent r e l a t i o n s h i p to the s i z e , t o t a l volume or growth rate of natural food p a r t i c l e s . This may be due to p e c u l i a r i t i e s i n the method of c a l c u l a t i o n of the index. Figure 10 shows that there i s no r e l a t i o n s h i p between e l e c t i v i t y values for any s i z e c l a s s and the t o t a l p a r t i c l e volume of any experiment. 148 Figure 11. Relationship between the elec subacuta on natural p a r t i c l e s (C 0/C.) i n each Coulter Count t i v i t y values (El) of Tintinnopsis and the changes i n c o n t r o l values er s i z e c l a s s . 149 o o ^ » CO -o t o 1^  coo -o oo — — K tv IT) o o co IT) CN o oooo to <*> ^ C N C N C N c o c o C N r o c o 00 A 1 CM CN CO CO co CN CN C N ^ C N d 150 Poulet (1973, 1974) has investigated the feeding rates of the small n e r i t i c copepod Pseudocalanus minutus on natural p a r t i c l e s with a Coulter Counter technique. Poulet also used Ivlev's e l e c t i v i t y index, and found that although p o s i t i v e and negative e l eettivity values were found i n a l l of h i s f i v e a r b i t r a r y s i z e classes from 1.5 ^im to 144 ^im, the frequency of p o s i t i v e values increased with the s i z e of the c l a s s to about 60 /lm diameter and then de-creased. E l e c t i v i t y values were mostly negative f o r Poulet's p a r t i c l e s be-tween 1.58 and 9 jam, except when p a r t i c l e s of t h i s s i z e reached 40 - 60% of the t o t a l concentration. Poulet (1974) found that food consumption i n P_. minutus was strongly correlated with the t o t a l p a r t i c l e volume i n a l l s i z e classes except i n the s i z e range below 3.57 u^m. Poulet therefore argued that P_. minutus has a very strong opportunistic feeding behaviour and a very ef-f i c i e n t u t i l i z a t i o n of the standing stock whatever i t s s i z e d i s t r i b u t i o n (and see General Discussion). Poulet's method of c a l c u l a t i n g feeding r a t e and standing stock and there-fore - e l e c t i v i t y i n d i c e s , comes under some suspicion p r i m a r i l y because he calculated feeding rate as merely the d i f f e r e n c e between f i n a l c o n t r o l and experimental p a r t i c l e concentrations. This means that he has used a l i n e a r method of c a l c u l a t i o n (see Methods Section) which may well be inappropriate f o r h i s p a r t i c l e concentrations and experimental periods and could lead to large over or under-estimation of the feeding r a t e . Nevertheless h i s con-clusions on the feeding 'strategies' of P_. minutus may well hold, but i t should be remembered that t h i s e l e c t i v i t y index indicates changes i n the proportion of items of a p a r t i c u l a r s i z e eaten, and does not n e c e s s a r i l y i n -dic a t e s e l e c t i o n by the predator. In the Coulter Counter experiments i n t h i s study Tintinnopsis subacuta does not appear to eat proportionately more of 151 a s i z e c l a s s of natural p a r t i c l e s when that s i z e c l a s s contains a large t o t a l volume of p a r t i c l e s (Figure 9), nor when the t o t a l volume i n a l l s i z e classes i s large (Figure 10). Therefore T_. subacuta does not appear to be as 'oppor-t u n i s t i c ' a feeder as P_. minutus; but not a l l the p a r t i c l e s i n a s i z e c l a s s may be equally a v a i l a b l e for predation nor equally edible (and see Section 4a). T_. subacuta l i k e P_. minutus eats p a r t i c l e s i n a l l s i z e classes i n i t s range but takes disproportionately more from the middle s i z e classes (3.57 to 7.12 ^m mean diameter), more frequently than from the p a r t i c l e s of l e s s than 3.57 ^ um or more than 7.12 jum diameter. These middle s i z e classes are those which show the greatest p o t e n t i a l growth rate i n these Coulter Counter experiments. The methods used by Poulet ( l o c . c i t . ) d i d not allow him to discuss the r e l a t i o n ^ ship between p o t e n t i a l growth rate and e l e c t i v i t y . " The e f f e c t of t i n t i n n i d feeding on the 'control' of phytoplankton populations I t i s important to t r y to a s c e r t a i n the conditions under which t i n t i n n i d s and other microzooplankton may 'control' phytoplankton populations composed of r e l a t i v e l y small organisms. Eigures 12,13 and 14 show the t o t a l e f f e c t of predator feeding i n these Coulter Counter experiments, and the number of predators/ml i n the same experiments. General 'control' i s presumed to have E C taken place when _2 < 1 and _2 ,> 1. A l l values have been recalculated from E l C l the data f o r an experimental duration of 24 hours, and the predator concen-t r a t i o n s -shown are those calculated to be alive/ml a f t e r 12 hours. The NTC values are given. The data are so v a r i a b l e that they support only the most tent a t i v e statements, but i t i s clear that natural concentrations of t i n t i n -nids ;(at l e a s t of T_. subacuta) , apparently under some circumstances, reduce the biomass of growing phytoplankton populations wi t h i n 24 hours, thus exer-t i n g a c o n t r o l l i n g influence. 152 Figure 12. Relationship between the changes i n the t o t a l p a r t i c l e volume of Coulter Counter control (C 2/C ) and experimental (E /E ) containers at various concentrations of t i n t i n n i d s per ml. '1.0' - Tintinnopsis subacuta on natural p a r t i c l e s . - T_. parvula on natural p a r t i c l e s . 153 o O i m m i to 6 — i 154 Figure 13. Relationship between the changes i n the t o t a l p a r t i c l e volume of Coulter Counter c o n t r o l (C^/C^) and experimental (E2/E^) containers at various concentrations of Tintinnbpsis^subacuta per ml. on laboratory food. 155 156 Figure 14. Relationship between the changes i n the t o t a l p a r t i c l e volume of Coulter Counter control (C^/C^) and experimental (E2/E^) containers at various concentrations of predators per ml. '1.0' - Stenosomella ventricosa on laboratory food. (lT(}) - Barnacle and copepod n a u p l i i on laboratory food. - Barnacle and copepod n a u p l i i on natural p a r t i c l e s . 158 A Coulter Counter experiment with R o t i f e r s Of several attempted Coulter Counter experiments with planktonic r o t i -f e r s , only one gave useful r e s u l t s . This was much shorter thantthe e a r l i e r experiments with t i n t i n n i d s and has been analysed separately and the r e s u l t s shown i n Table 29. Of the two species used: Synchaeta l i t t o r a l i s (about 500 x 200 ^im) i s the l a r g e s t marine planktonic r o t i f e r i n the study area; and the smaller species i s also parobablhy. of the genus Synchaeta and i s about 250 x 100 pm. i n s i z e . A l l animals used were females; many :S. l i t - t o r a l i s females with external eggs do not take food (personal observation), and at such times t h e i r eggs have a darker, rougher appearance than usual. The l a t t e r may be miotic (or f e r t i l i s e d ) eggs and are possibly among the l a s t produced by such a female. None of the S^. l i t t o r a l i s i n t h i s experiment appeared to carry m i c t i c eggs. In t h i s experiment, the same con t r o l was used f o r both r o t i f e r species, and the prey was D u n a l i e l l a t e r t i o l e c t a i n f i l t e r e d seawater. The duration of the experiment was 7.42 hours f o r S_. l i t t o r a l i s and 7.17 hours for Synch- aetaa sp., •' There were 0.65 S^. l i t t o r a l i s / m l and 0.60 Synchaeta sp./ml. The 'exponential' equation (see Methods Section) was used to c a l c u l a t e the feeding rate (FR/P). In Table 29 p o s i t i v e values of FR/P i n d i c a t e a net increase i n p a r t i c l e volume i n that s i z e c l a s s , and negative values indi c a t e a net l o s s ofi p a r t i c l e volume. The l a t t e r i s interpreted here as due to consumption by r o t i f e r s . Table 29 shows that p a r t i c l e consumption by both species occur-red in. s i z e classes of mean diameter 7.12 to 14.3 ;um; and that a net increase of p a r t i c l e s occurred i n s i z e classes 2.28 to 7.12 jxm. These opposite trends were weakest i n the middle s i z e c l a s s e s , and there seems to be a gradual change from production to consumption of p a r t i c l e s with increasing p a r t i c l e T A B L E 29. Results of Coulter. Counter experiment with Synchaeta littoralis and Synchaeta sp. eating Dunaliella tertiolecta. '• Mean Diameter of Size Class (um) 2.82 3.57 4.49 5.66 7.12 8.98 11.3 14.3 Total C a / C i 1.14 0.94 0.51 0.79 0.69 0.54 2.43 13.4 0.86 S. littoralis Ej. (um 3 x 103) 87.4 88.7 109.6 88.6 132.2 238.3 64.0 25.9 834.7 Ea (um 3 x 103) 152.7 98.9 98.0 87.7 80.7 120.6 42.6 13.0 694.2 Log mean E (Um 3 x 103) 117.1 93.7 103.5 88.2 104.4 172.7 52.6 19.0 751.2 F R W m 3 x 1 0 3 / h r / predator +10.4 . +3.3 +12.0 +4.1 -2.7 -2.3 -14.4 -12.8 N T C -2.2 ESO -32.1 F R ml/hr/predator -- -- -- 0.026 0.013 0.274 0.674 N T C 0.0029 ESO 0.0427 E i 93.8 89.0 110.0 Synchaeta sp. 87.5 1 50.7 305.0 64.6 17.3 918.0 Ea 195.5 100.8 98.1 83.4 103.1 '100.5 56.0 24.5 761.9 Log Mean E 139.0 95.3 103.9 85.8 128.5 184.2 60.3 20.7 .838.3 F R u m 3 x 1 0 3 / h r / predator +19.4 +4.1 +13.5 +3.7 - -0.2 -21.0 -14.4 -10.8 N T C -5.7 ESO -46.4 F R ml/hr/predator 0.002 10.114 0.239 0.522 N T C 0.0067 ESO 0.0553 160 s i z e . In t h i s r e l a t i v e l y short experiment, p a r t i c l e production at these sizes i s probably caused by the fragmentation of large p a r t i c l e s or by the egestion of o l d food and not by d i f f e r e n t i a l p a r t i c l e growth. The v a l -ues i n Table 28 ind i c a t e that values greater than 1.0 were associated with a high feeding r a t e . However, the o v e r a l l n O ^ / C ^ v a l u e w a s 0.86. As i n the other Coulter Counter experiments the ESO values were higher than the NTC values;.bbut i n t h i s case the l a t t e r were also negative. Feeding rates calculated as m l / h r / r o t i f e r were, for the t o t a l p a r t i c l e volume:0.0029 (NTC) and 0.0427(ESO) for S_. l i t t o r a l i s ; and 0.0067(NTC) and 0.0553(ES0) for Synchaeta sp.. Those s i z e classes which contained most of the D. t e r t i o - l e c t a were 7.12 to 11.3 jam diameter; and the feeding rates i n m l / h r / t i n t i n n i d f o r the t o t a l of the l a t t e r were: 0.0586 for S_. l i t t o r a l i s and 0.00954 for Synchaeta sp.. I t i s s u r p r i s i n g that the feeding rate of Synchaeta sp. on p_. t e r t i o l e c t a here exceeds that of the much larger S^ . l i t t o r a l i s . At these rates 100% of the p_. t e r t i o l e c t a would have been eaten i n 10 to 17 hours. The feeding rates shown i n Table 29 exceed those of any t i n t i n n i d species i n t h i s study by a f a c t o r of 2 (Coulter experiments) to 5 or more(accumu-l a t i o n experiments). R o t i f e r s i n inshore waters i n t h i s area may oc c a s i o n a l l y a t t a i n concentrations as high as 3 or 4/ml, and at such times an equivalent t o t a l feeding rate by t i n t i n n i d s would require concentrations of perhaps 15 to 20 T_. subacuta/ml. Concentrations such as t h i s of large t i n t i n n i d s r a r e l y occur. Although these two r o t i f e r species have f a s t e r feeding rates than t i n t i n n i d s and rapid reproductive rates; at times they do not feed at a l l , and they are much less common than t i n t i n n i d s i n cold or h i g h - s a l i n i t y water. 4 3 Tintinnopsis subacuta has a maximum c e l l volume of about 7 x 10 ; a large female _S_. l i t t o r a l i s i s about 100 times l a r g e r . Therefore, on the basis of 161 i n d i v i d u a l volume, the t i n t i n n i d i s a considerably more e f f e c t i v e predator than the r o t i f e r . V a r i a b i l i t y i n feeding rates i n Coulter Counter experiments The four most l i k e l y causes of v a r i a b i l i t y i n these experiments were: 1) v a r i a b i l i t y between r e p l i c a t e s of co n t r o l and experimental samples; 2) divergence between growth rates of prey items i n control and experimental samples; 3) i n v a l i d assumptions about mortality of the predators; 4) d i f -ferences i n the feeding response of predators to various prey types, p a r t i -c u l a r l y i n those experiments where one or a few prey types may have been dominant. V a r i a b i l i t y between c o n t r o l r e p l i c a t e s i s not r e a d i l y e x p l i c a b l e , but Oct c e r t a i n l y any i n i t i a l d i fferences would have been most enlarged i n the longest experiments. The combination of v a r i a b i l i t y i n control and experimental samples would be m u l t i p l i e d i n the calculationoof feeding r a t e s . The assum-ption that the p o t e n t i a l changes i n p a r t i c u l a t e volume i n each s i z e c l a s s i n the experimental vessel p a r a l l e l the changes i n those s i z e classes i n the con t r o l v e s s e l s , was also l e a s t v a l i d i n the longest experiments. Several of the experiments were too long, but i t was often d i f f i c u l t to obtain s i g n i f i -cant changes i n p a r t i c u l a t e volumes i n more optimal periods of time such as 6 to 8 hours, p a r t i c u l a r l y with low natural concentrations of t i n t i n n i d s . In order to c a l c u l a t e feeding rates the assumption was made that the t i n t i n n i d s which died during the course of the experiments did so at a u n i -form r a t e . This assumption was not v e r i f i e d . If most of the dead t i n t i n n i d s had died near the s t a r t , or conversely near the end of experiments; the use of the above assumption would greatly underestimate or overestimate .. 1 6 2 r e s p e c t i v e l y , the feeding rate of the remainder. In any future Coulter Coun-ter experiments with t i n t i n n i d s , the number, condition and accumulated food contents of t i n t i n n i d s should i d e a l l y be checked at i n t e r v a l s . As has been seen i n r e s u l t s of accumulation experiments (Section 4a ( i i ) ) , some t i n t i n n i d species show apparent d i f f e r e n t i a l predation and s e l e c t i o n on some types of food. This s e l e c t i o n usually takes a negative form i n that some food items are eaten disproportionately l e s s when they are presented together with another food item which i s apparently more e a s i l y caught even i n low concent-ra t i o n s . The q u a l i t a t i v e response of t i n t i n n i d s and other microzooplankton to various prey types should also be checked i f po s s i b l e l d u r i n g Coulter Coun-ter experiments on natural samples. 163 5) General Discussion Of the 13 species of t i n t i n n i d s mentioned i n t h i s study: much of the information has been gained from one common and r e l a t i v e l y large species, Tintinnopsis subacuta; somewhat l e s s has come from 4 or 5 other species of moderate s i z e and frequency; and s t i l l l e s s from 3 or 4 more species mostly found i n mid-summer. V i r t u a l l y no information has come from the 2 smallest, the 2 l a r g e s t , and one very rare species. The two smallest species Tintinnopsis nana and Tintinnopsis rapa did not eat enough v i s i b l e p a r t i c u l a t e material to give much information with the techniques used here. The l a r g e s t species, F a v e l l a serrata and Eutintinnus latus were too rare to be of much p r a c t i c a l use, although some information was obtained from them, e s p e c i a l l y from E. latus (Section 4b) . Of the other c i l i a t e microzooplankton encountered, the h o l o t r i c h s Prorodon sp. and Mesodinium rubrum were never seen to ingest p a r t i c l e s , a l -though the former was very occasionally seen to contain food material and may have a very rapid d i g e s t i o n rate. There i s one Coulter Counter experimental r e s u l t from the h o l o t r i c h T i a r i n a fusus. This species occasionally occurred i n large numbers but i s probably p r i m a r i l y a histophage. Of the o l i g o t r i c h c i l i a t e s which are c l o s e l y r e l a t e d to t i n t i n n i d s , several genera and species were observed during this study. The smallest species (15 to 30 ,um) , almost c e r t a i n l y ingested b a c t e r i a and/or dissolved organic material; those of a s i m i l a r s i z e range to t i n t i n n i d s appeared to contain s i m i l a r food to t i n -t i n n i d s i n s i m i l a r amounts, and were often very numerous; and the l a r g e s t (200-300 um) were predatory, l a r g e l y uponntintinnids. Some of the non-tin-t i n n i d c i l i a t e s may be 'quasi-symbiotic' at times (Blackbourn et.al_, 1973) but t h i s does not appear to be true of t i n t i n n i d s . 164 One Coulter Counter experiment was c a r r i e d out with two species of r o t -i f e r s , and t h e i r feeding rates were about 2 to 5 or more times as f a s t as those of Tintinnopsis subacuta. (Table 30). The feeding rates of mixtures of barnacle n a u p l i i and small copepod n a u p l i i ( c h i e f l y the former) when measured with the Coulter Counter method, were very s i m i l a r to those of the two t i n t i n n i d species used i n s i m i l a r experiments. I t seems l i k e l y from t h i s fragmentary information that the p o t e n t i a l o v e r a l l feeding e f f e c t of t i n -t i n n i d s on natural populations of phytoplankton, i s a a t l e a s t as great as that of any other group of microzooplankton used i n t h i s study. It i s possible that the o v e r a l l e f f e c t of the feeding of o l i g o t r i c h s i s greater than that of t i n t i n n i d s , but there i s no quantitative information on the feeding r a t e s of the former. O l i g o t r i c h populations seemed to be even more transient and v a r i a b l e than those of t i n t i n n i d s , and the population feeding e f f e c t of either group would be extremely d i f f i c u l t to evaluate over periods of more than a day or two. In comparing the r e s u l t s of t h i s study with those of other authors, the great v a r i a b i l i t y which has b e d e v i l l e d the project makes any but the most general statements untenable. However, the following comments on some d e t a i l s and some general trends are probably j u s t i f i e d . There are no other quanti-t a t i v e data on t i n t i n n i d or o l i g o t r i c h c i l i a t e feeding rates with which to compare t h i s work, but the feeding rates of some f r e e - l i v i n g semi-planktonic c i l i a t e s have been examined i n the papers by Goulder (1972), Hamilton and Preslan (1969) and Pavlovskaya (1973). Rapport e t . a l . (1972). have discussed the food preferences of the s e s s i l e c i l i a t e species Stentor coeruleus, and Berger (1971), Curds and Cockburn (1971) and Ricketts (1971, 1973) and others have discussed feeding i n the freshwater bacteria-eating species Paramecium  a u r e l i a and Tetrahymena pyri f o r m i s . Most authors who have worked with 165 protozoa have used methods d i f f e r e n t from those used i n t h i s study, e.g. rad i o a c t i v e tracers (Pavlovskaya, 1973); the estimation of weight or t o t a l c e l l y i e l d , etc. (Curds and Cockburn, 1971; Hamilton and Preslan, 1969) or extrapolations from rates of l o s s to feeding rates (Goulder 1972) . Berger (1971) made counts of food vacuoles l a b e l l e d with radioactive b a c t e r i a , and Ricketts (1971) counted food vacuoles which contained l a t e x p a r t i c l e s . Rapport et^.al_. (1972) counted the accumulations of food' i n Stentor coeruleus a f t e r 1 hour. T i n t i n n i d s eat almost anything of l e s s than a c e r t a i n s i z e (Table; 2), but d i f f e r e n t i a l predation does occur i n some species on some prey' types (Section 4a). For example,' Tintinnopsis subacuta eats disproportionately more E u t r e p t i e l l a sp. than other prey species i n mixtures; and t h i s appears to depress the ing e s t i o n of the other species compared to controls when the l a t t e r are presented si n g l y to T_. subacuta. This might be considered as negative s e l e c t i o n , but i f de l i b e r a t e , i t s u t i l i t y i s not obvious. On the other hand, Cryptomonas sp. and Isoselmis sp. (Cryptophyceae) are eaten l e s s than proportionately by T_- subacuta. T_. cylindrica,- T_; parvula and Stenosomella ventricosa when mixed with other prey items, and very l i t t l e at any time. True negative s e l e c t i o n by these t i n t i n n i d s on Cryptomonads was also found i n some experiments (Section 4a), On the other hand Tintinnidium mucicola accumulates a l l food items at a generally slower rate than the above t i n t i n n i d species, and has an apparently disproportionately negative feeding r e a c t i o n to most prey species i n mixtures but to Isoselmis sp. and other Cryptomonads. The feeding rate of S_. ventricosa and Tintinnopsis c y l i n d r i c a on most small prey items (such as Monochrysis l u t h e r i ) i s f a i r l y s i m i l a r to that of T. subacuta to which they are s i m i l a r i n s i z e . However, S_. ventricosa eats much l e s s E u t r e p t i e l l a sp. from_a s i m i l a r concentration of c e l l s , than 166 does T_. subacuta, and T_. c y l i n d r i c a has never eaten E u t r e p t i e l l a sp. i n the laboratory although c e l l s which resemble euglenoids have been seen i n s i d e t h i s t i n t i n n i d species from f i e l d samples. I t has been shown that Eutintinnus  la t u s which i s larger than T_. subacuta, cannot subdue normal E u t r e p t i e l l a sp. but can eat the larger and slower prey species Cryptomonas profunda. (Section 4c). 12. latus does not seem to habituate to E u t r e p t i e l l a sp. and never f a i l s to attempt to ingest i t . T i n t i n n i d s may deal d i f f e r e n t l y with r e l a t i v e l y small prey. This kind of d i f f e r e n t i a l and sometimes negatively s e l e c t i v e predation has not been i d e n t i f i e d i n other groups of c i l i a t e s , but no doubt i t occurs. In contrast, Rapport (Rapport, elt.aJU 1972; and unpublished data) has found d e f i n i t e but t r a n s i t i v e p o s i t i v e feeding preferences i n Stentor coeruleus. S_. coeruleus,genera-My_y prefers large prey to small prey and' c i l i a t e s are preferred to f l a g e l l a t e s ; but the degree of preference changes with changes i n the absolute and r e l a t i v e concentrations of prey' types i n mixed samples. JL" coeruleus s h i f t s i t s preference i n the d i r e c t i o n favouring that;>prey species of r e l a t i v e l y high abundance. Feeding experiments with t i n t i n n i d s over a wide range of concentrations of mixed prey were not possible with the techniques used i n t h i s study; but i t i s possible that the degree of prey s e l e c t i o n seen i n Section 4a may change with changes i n the concentration of prey. I t i s important to note that whatever i t s e f f e c t on the mixed-prey populations, a preference for the most abundant prey item w i l l probably be of l e s s use to a predator than a preference f o r that ( e a s i l y digested) prey with the largest t o t a l biomass/ml (see comparative discussion of Coulter Counter experiments below). The feeding response of microzooplankton to prey may greatly depend upon the q u a l i t y of the prey, which may vary between prey species and c e r t a i n l y between d i f f e r e n t growth stages of the same species. 167 Such q u a l i t a t i v e d ifferences may explain some of the v a r i a b i l i t y seen i n t h i s study and i n the r e s u l t s of Rapport (unpublished data) and Strathmann (1971). However, several d i f f e r e n t species of phytoplankton have s i m i l a r proportions of the major n u t r i t i o n a l constituents, and when i n the exponental growth phase (as i n 'blooms' or from new laboratory cultures) should be simi-l a r l y useful to a predator. Predators must respond to the behaviour or surface c h a r a c t e r i s t i c s of prey, and therefore s e l e c t i o n cannot be based upon i t s n u t r i t i o n a l composition per se. Also i t 1 i s d i f f i c u l t to understand the b i o l o g i c a l basis f o r the persistence of predation on a less - p r e f e r r e d prey type at very high concentrations of mixtures of prey types ( t h i s study Section 4a and Rapport, unpublished data), since most i n d i v i d u a l s of a l l prey species w i l l be rejected under such circumstances. The feeding rates of t i n t i n n i d s i n t h i s study i n general increase with the increase of c e l l volume among predator species. Also the search rates of various t i n t i n n i d species as seen i n Section 4b on items which can be ingested by a l l , do not vary by more than a factor of 5 or 10, and the range of c e l l volumes among these t i n t i n n i d s i s also approximately 10-fold. A feeding rate of 0.0042 m l / h r / t i n t i n n i d (Table 15) from E u t r e p t i e l l a sp. at 1,400 c e l l s / m l amounts to about 6 prey c e l l s ingested/hour, and i s equivalent 3 to about 3,000 11m / h r / t i n t i n n i d . To obtain t h i s volume of food T_. subacuta would have to ingest about 60 Monochrysis lutheri/hour or about 15 D u n a l i e l l a t e r t i o l e c t a / h o u r . At a feeding r a t e of 0.0042 m l / h r / t i n t i n n i d t h i s would require c e l l concentrations of 10 timescor 221^2 times that of E u t r e p t i e l l a  sp. f o r M. l u t h e r i and p_. t e r t i o l e c t a r e s p e c t i v e l y . From the r e l a t i o n s h i p s between temperature, s i z e , and reproductive rate shown for benthic c i l i a t e s by Fenchel (1968), i t can be estimated that the maximum reproductive r a t e of a t i n t i n n i d such as T_. subacuta might be about one d i v i s i o n per 37 hours 168 o 0 at 10 C and one per 15 hours at 20 C, i f i t could t o l e r a t e such a temperature fo r long. However, Gold (1971, 1973) has shown that Tintinnopsis heroidea (probably synonymous with T. subacuta) divides about once every 19 to 27 hours at 12.5°C i n continuous c u l t u r e . A rate of one d i v i s i o n i n 24 hours would give a maximum s p e c i f i c growth rate of 0.029 hr f o r a newly divided A 3 T_. subacuta (*4 x 10 /um ), At a growth/consumption r a t i o (K^) of 0.3 (Klekowski e t . a l . , 1972; PavMvskaya, 1973) t h i s would require a continuous 3 intake of 3,880 pm /hr or 9.7% t i n t i n n i d c e l l volume/hr or about 8 E u t r e p t i e l l a sp. or 80 Monochrysis l u t h e r i / h r . At a feeding rate of 0.0042 ml/hr, the prey c e l l concentration that o would be necessary for the maximum growth rate of a new T_. subacuta at 12.5 C would be about 1,900/ml E u t r e p t i e l l a , or about 19,000/ml M. l u t h e r i (0.95 ppm). Thus i t seems p l a u s i b l e that T_. subacuta could obtain enough food from the concentration of E u t r e p t i e l l a sp. i n Table 15 (0.75 ppm) to reach a repro-ductive rate of about 3/4 of i t s t h e o r e t i c a l maximum. Blooms of T_. subacuta and euglenoid and other large f l a g e l l a t e s , and temperatures of about 12 C a l l occur t y p i c a l l y i n l a t e spring i n t h i s area. However, i t i s almost c e r t a i n that feeding by T_. subacuta i s not continuous; and j u s t as the r e l a t i o n s h i p between feeding rate and digestion r a t e i s not a simple one, the r e l a t i o n s h i p between dig e s t i o n and growth may also be complex. Pavlovskaya (1973) gives f a i r l y simple models for the r e l a t i o n s h i p be-tween food'concentration, feeding r a t e on one type of food and reproductive r a t e s , f o r three species of c i l i a t e s . Her data has some'unacknowledged incon-s i s t e n c i e s , and i t i s not possible to judge the accuracy of the estimates of feeding rate. She gives the r e l a t i o n s h i p between feeding rate ( r a t i o n ) , maximum r a t i o n , and food concentration as: 169 R = Rmax ( l - e p ( k - k o ) ) (5) where R = r a t i o n , and p and k are constants. The r e l a t i o n s h i p between reproductive r a t e and r a t i o n i s given by: -g = a R"b C6) where g = time taken for a c i l i a t e to double i t s population s i z e (in days) and a and b are constants and are derived from the data. The values of R and g given by Pavlovskaya (loc. c i t . ) are as follows: -Keronopsis rubra , Q . n -5 l V " RGng/hr) = 5 x l 0 - 6 ( l - e - 8 0 ( k - ° - 8 x 1 0 *) (wet weight 10 mg) g ( d a y g ) = ^ ( R ) -0.215 Uroleptopsis v i r i d i s R(mg/hr) = 1 4 . 9 x l 0 ~ 5 ( l - e ~ 8 5 0 ( k ~ 0 * 2 x l ° g(days) = 0.205 (R) - 0 - 1 6 9 Condylostoma magnum _3 (wet weight 7x10 mg) R(mg/hr) = 4.9xl0- 4(l-e-°- 5 ( k-°- 5 )) Condylostoma magnum i s a very large free swimming c i l i a t e here feeding on dinoflagellates,and the other two species are bottom feeders on diatoms and ba c t e r i a . The maximum rations shown by Pavlovskaya (loc. c i t ) were obtained at food concentrations equivalent to 5% of c i l i a t e body weight/hr at 2,000 2 diatoms/Cm for Keronopsis rubra; and equivalent to 6% of body weight/hr fo r Condylostoma magnum. The f a s t e s t rates were about 3 times f a s t e r than, and occurred at food concentrations of about 1/3 of, the l a t t e r . As Tintinnopsis subacuta i s somewhat smaller than these two species, i t seems reasonable that i t s feeding rate i n terms of percentage of c e l l volume/hr as calculated above i s a l i t t l e higher than that of Keronopsis rubra and Condylostoma magnum. Hamilton and Preslan (1969) showed that although the maximum s p e c i f i c 170 growth rate of the small c i l i a t e Uronema marinum i s very high (0.147 h r ) ; i t feeds s u f f i c i e n t l y slowly that the concentration of bacteria necessary for maximum growth (0.49 ug C/ml or 10 ppm) would be very r a r e l y found i n the sea, except perhaps on the corpses of planktonic organisms. Therefore, they termed U. marinum an 'opportunistic' predator. I t i s l i k e l y that very small t i n t i n n i d s such as Tintinnopsis nana and small o l i g o t r i c h c i l i a t e s have s i m i l a r trophic c h a r a c t e r i s t i c s . An approximate estimation of the food required for maximum c e l l growth i n T_. nana s i m i l a r to that made f o r T_. subacuta (above) gives a value of about 60 x 10 Monochrysis l u t h e r i / m l , or even more of a smaller prey. This i s equivalent to about' 3.0 ppm by volume, a l e v e l so high as to be r a r e l y found among p a r t i c l e s of l e s s than 5 /im d i a -meter i n t h i s area. T_. nana has very transient 'blooms' i n t h i s area, i n which i t may become as abundant as 15 to 20/ml. Hamilton and Preslan (1969) also found that IJ. marinum underwent up to 20-fold changes i n c e l l volume, which may be a consequence of a time lag i n the rate of d i v i s i o n . Canale e_t ,al_. (1973) noticed a s i m i l a r phenomenon with Tetrahymena p y r i f ormis. There was no evidence of great v a r i a t i o n i n t i n t i n n i d c e l l volumes i n t h i s study. There has been considerable discussion of ' l o s s ' rates and of the r e l a -tionship between hunger, s t a r v a t i o n , and the feeding rates of t i n t i n n i d s i n t h i s section on accumulation experiments. No f i r m conclusions could be drawn from t h i s discussion. Berger (1971) found that the l o s s r a t e of food vacuoles i n feeding Paramecium a u r e l i a was an exponential function; and Goulder (1972) found the same r e s u l t at a much slower rate i n starved i n d i v i d u a l s of the very large c i l i a t e Loxodes magnus. Goulder ( l o c . c i t . ) equated l o s s rate with feeding rate and estimated the feeding rates of L_. magnus by extrapolating the rate of food l o s s back to zero time. However, this produces a very low 171 feeding rate f o r such a large c i l i a t e ; and as there i s some evidence that a feeding c i l i a t e loses food even i n i t i a l l y at a f a s t e r r a t e than a starved c i l i a t e (Rapport, unpublished data; t h i s study Section 4a), t h i s method of c a l c u l a t i o n may be suspect. Canale, e t . a l . (1973) and Gold (1971) have found that s t a r v i n g c i l i a t e s s u f f e r r a p i d m o r t a l i t y and c e l l l y s i s i n laboratory culture; and i n t h i s study i t was found that the feeding a b i l i t i e s of t i n -t i n n i d s were considerably reduced a f t e r complete sta r v a t i o n f o r about 30 hours, which may i n d i c a t e moribundity. More data i s needed on t h i s subject. The only other behavioural study of feeding i n c i l i a t e d microzooplankton was that of Strathmann, (1971). Strathmann studied the d e t a i l e d feeding behaviour and numbers of food c e l l s accumulated by the planktonic larvae of 15 species of echinoderms. He found that these larvae regulate t h e i r feeding r a t e on p a r t i c l e s with a v a r i e t y of methods. Echinoderm larvae show spon-taneous v a r i a b i l i t y of feeding rates over short periods of time even i n f i l t e r e d water. Likewise the v a r i a b l e response of i n d i v i d u a l t i n t i n n i d s to two apparently s i m i l a r successive p a r t i c l e s may spring from some purely i n -ternal p h y s i o l o g i c a l changes. P a r t i c l e s are ingested by echinoderm larvae by l o c a l induced reversals of beating by • some c i l i a of the c i l i a r y band, without the use of mucus. Feeding may be stopped or reduced by passing par-t i c l e s over the c i l i a t e d band with the water. Strathmann e t . a l . (1972) state that mucus i s probably used by many f i l t e r feeders only i n the r e j e c t i o n or egestion of p a r t i c l e s and not i n the process of ingestion. The d e t a i l s of the ingestion of very small p a r t i c l e s by t i n t i n n i d s could not be seen i n t h i s study although the o r a l plug may play some part i n the process (Section 4c). Echinoderm larvae can sort p a r t i c l e s i n s i d e the gut; and i f necessary expel them by muscular contractions, or i f they are i n d i g e s t i b l e sort them to bypass the stomach towards the-anus. It i s not known i f t i n t i n n i d s can 172 s o r t p a r t i c l e s i n t e r n a l l y . P a r t i c l e s are rejected before reaching the gut by the larvae stopping or reversing the beat of a l l c i l i a i n the c i l i a r y band, and t h i s seems analogous to the r e j e c t i o n behaviour of t i n t i n n i d s (Section 4b). Strathmann (1971) found that the rate of passage of food through the gut i n echihoderm larvae was extremely i r r e g u l a r , but generally increased with an increase i n the ingestion rate. This observation may be analogous to the 'forcing e f f e c t ' of new food on the r a t e of l o s s of o l d food by t i n -t i n n i d s seen i n the accumulation experiments i n t h i s study (Section 4a). Echinoderm larvae can ingest p a r t i c l e s ( e s p e c i a l l y 'discs' or 'spheres') of a s i z e of up to 75 to 100% of the diameter of the oesophagus or 80 to 100 jum i n most species. Ti n t i n n i d s can ingest p a r t i c l e s f a r larger than the undis-tended diameter of t h e i r cytopharynx and carnivorous c i l i a t e s are even more impressive i n this regard. Planktonic Crustacea r a r e l y ingest p a r t i c l e s which are r e l a t i v e l y as large as t h i s . Most echinoderm larvae can ingest large t h i n diatom c e l l s (200 x 30 jim) but l i k e t i n t i n n i d s they cannot deal with long chains of c e l l s of any s i z e . Strathmann (loc. c i t ) ) found that although there was no p o s i t i v e s e l e c t i o n of or preference f o r , food items of various types by echinoderm larvae there was some d i f f e r e n t i a l ease of capture; and he thought that the ingestion of very large items 'interfered' with the ingestion of smaller items (apparent negative s e l e c t i o n ) , though not v i c e versa. D i f f e r e n t i a l predation and apparent negative s e l e c t i o n i n mixed-prey s i t u a t i o n s was seen i n t i n t i n n i d s i n t h i s study (Section 4a); but any 'interference' by one food item with another i s u n l i k e l y to be a mechanical process at l e a s t i n t i n t i n n i d s , due to the time l a g between ingestion events even at high food concentrations. 173 Strathmann (1971) ca l c u l a t e d clearance (feeding) rates i n two ways which were e s s e n t i a l l y s i m i l a r to those used i n t h i s study i n Sections 4a and 4b, except that the periods of accumulation were shorter i n the more rapidly-feeding larvae than i n t i n t i n n i d s . Rates determined by^direct obser-vation of echinoderm larvae were higher and more v a r i a b l e than those deter-mined from food accumulation. Very l i t t l e ingestion was observed by t i n t i n -nids i n t h i s study; but the 'observed' feeding rate of Tintinnidium mucicola (Table 26) was somewhat greater than feeding rates calculated f o r t h i s species by the accumulation method. As there can be r e j e c t i o n of p a r t i c l e s from the mouth of echinoderm larvae,.and as observations were made over very short periods of time i n both kinds of organisms; t h i s discrepancy i n feeding rates i s not too s u r p r i s i n g . Echinoderm larvae do not feed at a rate s u f f i -c i e n t to pack the gut with algae f o r long periods of time. The optimal food concentrations (OFC) for echinoderm larvae are lower, and feeding rates decline more r a p i d l y at high food concentrations than i n some la r g e r types of microzooplankters (e.g. see Table 30). Although l i t t l e r e l i a b l e information on OFC values has been obtained during t h i s study: from some of the r e s u l t s i n the Section on accumulation experiments (Section 4a), i t seems that i n t i n t i n n i d s also (e.g. i'n Tintinnopsis subacuta) OFC l e v e l s may be as low as 1,500 c e l l s / m l of E u t r e p t i e l l a sp. or more of smaller prey. The feeding rates found i n 7 species of echinoderm larvae varied with food, species and growth phase between 0.02 and 0.53 ml/hr/larva. These rates are 2 to 50 times greater than the feeding rate of Tintinnopsis subacuta (Table 30). However t i n t i n n i d s are f a r more than 2 to 50 times more abundant than echinoderm larvae or any organism of s i m i l a r s i z e , i n t h i s area. Some of the feeding p p e c u l i a r i t i e s of the t i n t i n n i d s i n t h i s study 174 c o r r e l a t e well with q u a l i t a t i v e evidence from f i e l d samples. For example, r e l a t i v e l y high natural concentrations of euglenoid f l a g e l l a t e s are almost always accompanied by large numbers of a c t i v e Tintinnopsis subacuta, many of which w i l l be undergoing d i v i s i o n . R e l a t i v e l y large numbers of cryptomonad f l a g e l l a t e s are usually accompanied by r e l a t i v e l y large numbers of Tintinnidium  mucicola. F a v e l l a serrata i s at i t s most abundant l o c a l l y when there are f a i r l y high numbers of d i n o f l a g e l l a t e s i n f i e l d samples i n l a t e summer or f a l l . There i s some q u a l i t a t i v e evidence from studies of toxic 'red-tides' i n Eastern Canada f or the predation of Fav e l l a sp. on large d i n o f l a g e l l a t e s . There are few data on the e f f e c t of the feeding of c i l i a t e s or other microzooplankters on natural populations of prey organisms. Goulder (1972) estimated that the fresh-water c i l i a t e Loxodes magnus would remove a neg-l i g i b l e f r a c t i o n of the dominant a l g a l food items i n one day. His methods of c a l c u l a t i o n may have l e d to an under-estimate. Most of the other i n f o r -mation comes from the work of Parsons and LeBrasseur (1970) , Parsons e t . a l . (1967) and Poulet (1973, 1974) on the feeding rates and food webs of n e r i t i c planktonic Crustacea. These authors u t i l i s e d Coulter Counter techniques and some of t h e i r r e s u l t s have been discussed i n that part of the study (Section 4c). The feeding rates of Tintinnopsis subacuta and Stenosomella ventricosa have been measured with the Coulter Counter technique to vary between about 0.33 and 3.8% m l / h r / t i n t i n n i d depending upon the method of c a l c u l a t i o n . The o v e r a l l e f f e c t of t i n t i n n i d s on 'populations' of natural p a r t i c l e s ( l i v i n g or i n e r t ) i s sometimes to increase the volume of p a r t i c l e s i n some size c l a s s e s , or o v e r a l l . Similar r e s u l t s were found i n experiments with two species of r o t i f e r s i n t h i s study, and by Poulet (1973) on Pseudocalanus 175 minutus. Pseudocalanus minutus appears to eat p a r t i c l e s from 3 to 100 -114 pm. diameter, but apparently only eats those smaller than 9 um when they are r e l a t i v e l y very abundant; and i n general may eat the greatest volume/ hour from those si z e classes which contain the greatest t o t a l volume. This i s also seen i n some of the r e s u l t s of Parsons e t . a l . (1967). The s i t u a t i o n i n Tintinnopsis subacuta i s much l e s s c l e a r - c u t . Natural p a r t i c l e s as small as 1 pro. are eaten, but much l e s s i n proportion to t h e i r t o t a l volume than par-t i c l e s from 3 to 8 ^ im diameter. This apparent d i f f e r e n t i a l s i z e predation i n the t i n t i n n i d i s confused by the f a c t that p a r t i c l e s of the size that i s most c o n s i s t e n t l y eaten are also those with the highest p o t e n t i a l rates of increase (Section 4c). However, i n those experiments showing the greatest feeding rates of T_. subacuta on natural samples, and also the most co n s i s t e n t l y p o s i -t i v e e l e c t i v i t y values, the dominant prey item was a species of E u t r e p t i e l l a . T_. subacuta showed the same c h a r a c t e r i s t i c s i n the r e s u l t s of the accumulation experiments (Section 4a). In a d d i t i o n to d i f f e r e n t i a l predation on various sizes of p a r t i c l e s , several authors (Adams and Steele, 1966; Parsons, et^.aJL. 1967) have estimated that f i l t e r - f e e d i n g by planktonic crustacea may stop i n very low t o t a l con-centrations of food. This phenomenon was also judged to be present by extra-polations from Coulter Counter data i n Tg. .subacuta and S_. ventricosa f o r which i t i s probably not u s e f u l , but these extrapolated lower threshold values were extremely low i n these two species. Bearing i n mind the behaviour of t i n t i n n i d s , the use of such extrapolations i s probably not j u s t i f i e d i n t h i s case. Parsons e t . a l . (1967) and Poulet (1973, 1974) have demonstrated that the p o t e n t i a l growth of natural phytoplankton populations can be co n t r o l l e d 176 i n some cases by natural concentrations of small planktonic crustacea. Such p o t e n t i a l c o n t r o l has also been demonstrated by Tintinnopsis subacuta i n some experiments i n t h i s study (Section 4c). However, the great v a r i a b i l i t y i n these r e s u l t s do not r e a d i l y allow the i d e n t i f i c a t i o n of the necessary conditions (e.g. the c e l l concentration of T_. subacuta) f o r such c o n t r o l . Approximate r e l a t i v e s i z e s , range of food s i z e , and maximum feeding rates are compared i n Table 30 for four very d i f f e r e n t types of marine micro-zooplankton. Of the three metazoan organisms, only the r o t i f e r i s numerous enough l o c a l l y to have a comparable population feeding e f f e c t to the t i n t i n n i d over the s i z e range of food common to both, despite the greater feeding rate of the i n d i v i d u a l r o t i f e r . C e r t a i n l y manyyother large and small zooplankton and benthic f i l t e r feeders w i l l also a f f e c t phytoplankton populations i n th i s and other coastal areas. However i t seems highly l i k e l y that c i l i a t e s w i l l have a greater e f f e c t on the biomass and d i v e r s i t y of the productive phyto-plankton of l e s s than 10 pm diameter than any other zooplankton organisms. Due to considerable food s i z e overlaps, the r e l a t i v e p r o d u c t i v i t y and feeding impact of t i n t i n n i d and o l i g o t r i c h species on small natural p a r t i c l e s w i l l p a r t l y depend on the prey species and biomass d i s t r i b u t i o n . For example, the maximum reproductive rate of Tintinnopsis nana may be 4 or 5 times that of T_. subacuta; and at such a rate T_. nana would soon be so numerous as to have a greater feeding e f f e c t than larger t i n t i n n i d s on p a r t i c l e s of 5 jim diameter. However, i n order to reach the maximum ra t e , T_. nana might require 4 to 5 times as many p a r t i c l e s of that si z e as would J_. subacuta, because of the f a s t e r search rate of the l a t t e r . T_. subacuta can also eat p a r t i c l e s 60 times larger than i s possible for T_. nana including T_. nana i t s e l f . Thus T_. subacuta may be able to reproduce at a rate equal to or greater than that of T A B L E 30. Approximate relative sizes and feeding rates of various types of marine microzooplankton. Organism Basis of Size Range Maximum FR Relative Size of food as percentage Size Comparison (spheres) (Um) ml/hr/pred Optimal Food ! Concentration (Total) um 3x 10a/ml (ppm) Maximum FR as per-centage body vol. or wt/hr Reference Tintinnopsis  subacuta (protozoan) Synchaeta littoralis (rotifer) Stronqylocentrotus dro e bac hiensis (echinoderm larva) (7x10* urn") 100 500 Estimated volume Length^ 2 1,-20 <5-7 <8 - 85 Av. 1.0 4.3 9.6 0.7 10.0 0.6 0.01 (est.) Present Study Present Study Strathmann (1971) ^seudocalanus  minutus (copepod) 1000 U g . C 100 17.0 15 2.3 Parsons and Lebrasseur (1970) and Poulet (1974) 178 T_. nana i n many natural s i t u a t i o n s . The same s i m p l i s t i c reasoning may apply equally w e l l to comparisons between other species of t i n t i n n i d s . The importance of t i n t i n n i d s as p o t e n t i a l ' c o n t r o l l e r s ' , competitors, or valuable prey i s s t i l l i i n doubt; but c e r t a i n l y the larger species, par-t i c u l a r l y T intinnopsis subacuta, eat at a s u f f i c i e n t r a t e and are numerous enough at times i n English Bay to f i l l a l l these categories. 179 (6) SUMMARY 1) General aspects of the feeding biology of 13 l o c a l species of t i n -t i n n i d s and some other microzooplankton were examined q u a l i t a t i v e l y and qu a n t i t a t i v e l y . 2) Ti n t i n n i d s and other microzooplankton ate a wide v a r i e t y of items: l i v i n g and i n e r t , natural arid unnatural, including other t i n t i n n i d s . 3) The maximum volume of food eaten was a function of the t i n t i n n i d c e l l volume taken over a l l species. T i n t i n n i d species of s i m i l a r volume were d i s s i m i l a r iri the maximum s i z e of t h e i r food. 4) There i s apparently no minimum s i z e of food f o r t i n t i n n i d s and a l l the l o c a l species ate bac t e r i a and other p a r t i c l e s of 1.5 um diameter or l e s s . 5) Several t i n t i n n i d species showed d i f f e r e n t i a l predation on laboratory cultures of phytoplankton. In various species these differences were based upon: the handling a b i l i t y of the predator; prey s i z e ; or prey type. 6) Also some types of laboratory phytoplankton were selected le s s than others from some mixed-prey si t u a t i o n s p a r t i c u l a r l y by Tintinnopsis subacuta. 7) The rates of accumulation of prey by four species of t i n t i n n i d were l i t t l e a f f e c t e d by temperature, but there was some evidence of a fa s t e r rate of l o s s of ingested food at very high temperatures. 8) The r e l a t i o n s h i p between the gain of new food and the loss of old food i n individual'! Tintinnopsis subacuta and Stenosbmella ventricosa was highly v a r i a b l e and may be heavily dependent upon the p h y s i o l o g i c a l h i s t o r y of the i n d i v i d u a l c e l l . The rate of gain of new food seemed to be l a r g e l y inde-pendent of the amount of old food i n a t i n t i n n i d , but the rate of loss of old food was on average f a s t e r i n those c e l l s which were gaining the greatest amounts of new food. 180 9) The feeding rates of a t i n t i n n i d species were extremely v a r i a b l e within and between experiments. Much of .this v a r i a b i l i t y was due to the use of t i n t i n n i d s from f i e l d samples and much was due to the experimental methods used. 10) Feeding rates as estimated by the three methods used showed f a i r q u a l i t a t i v e agreement but poor quantitative agreement. The feeding rate of Tintinnopsis subacuta i n accumulation experiments was the equivalent of 0.65% m l / h r / t i n t i n n i d or usually much l e s s . In Coulter Counter experiments T_. subacuta ate the average equivalent of 2.0% (minimum) or 3.8% (maximum) ml / h r / t i n t i n n i d on natural samples, and 0.33% (minimum) or 2.0% (maximum) ml / h r / t i n t i n n i d from cultures of laboratory phytoplankton. 11) The feeding rates of Tintinnopsis subacuta and Stenosomella ventricosa were p o s i t i v e l y correlated most co n s i s t e n t l y with the i n i t i a l and estimated mean t o t a l experimental p a r t i c l e volume. C o r r e l a t i o n c o e f f i c i e n t s between feeding rates and another seven v a r i a b l e s showed l a r g e l y i n e x p l i c a b l e trends. 12) I t has been shown that p r e d i c t i o n of t i n t i n n i d feeding rates from only a knowledge of the s i z e d i s t r i b u t i o n of p a r t i c l e biomass i n a natural sample would be impossible. 13) Feeding rates of T_. subacuta and S_. ventricosa i n Coulter Counter ex-periments showed no apparent upper asymptote below 0.76 ppm on natural samples and 1.92 ppm on laboratory cultures. However, i n accumulation ex-periments T_. subacuta showed apparent asymptotes (OFC) at 0.35 ppm on D u n a l i e l l a t e r t i o l e c t a and at 0.70 ppm on E u t r e p t i e l l a sp. These differences were not resolved. 181 14) The Ivlev e l e c t i v i t y indices of Tintinnopsis subacuta on natural samples were most c o n s i s t e n t l y p o s i t i v e i n the middle s i z e classes (3 to 7.5 um dia.) of i t s food range. These s i z e classes showed the greatest growth i n c o n t r o l s . The magnitude of the e l e c t i v i t y indices of t i n t i n n i d s on any prey type i n accumulation experiments p a r t l y depended on the amount of other prey types eaten. 15) Natural concentrations of Tintinnopsis subacuta can apparently control the growth of natural populations of phytoplankton l e s s than 20 um d i a . under some conditions; The most l i k e l y concentrations of T_. subacuta neces-sary f o r such control are unknown due to the great v a r i a b i l i t y of r e s u l t s . 16) An average feeding rate of T_. subacuta was compared with those of l a r g e r microzooplankters. 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Comparison avec d'autres Tintinnides et avec l e s autres ordres de c i l i e ' s . P r o t i s t o l o g i c a 8_: 369-386. (1973). Cortex et Perilemme de Cy t t a r o c v c l i s brandti ( c i l i e , T i n t i n n i d e ) . Remarques sur l e s structures c o r t i c a l e s des c i l i e s . J . C e l l B i o l . 59: 246. LeBrasseur, R.J. and O.D. Kennedy (1972). Microzooplankton i n coastal and oceanic areas of the P a c i f i c subartic water mass: a preliminary report, pp. 355-365 In ' B i o l o g i c a l Oceanography of the Northern North P a c i f i c Ocean'. Ed. by A.Y. Takenouti et. a l . Idemitsu Shoten, Japan. L u c k i n b i l l , L.S. (1973). Coexistence i n laboratory populations of Paramecium  a u r e l i a and i t s predator Didinium nasutum. Ecology 54: 1320-1327. Marshall, S.M. (1969). Protozoa - O r d e r : T i n t i n n i d a . Fiches d ' i d e n t i f i -c a t i o n du zooplankton. Cons. Perm. Int. pour L'exploration de l a Mer. Sheets 117-127. (1973). Respiration and feeding i n copepods. Adv. Mar. B i o l . I i : 57-120. Mitchison, J . (1971). The biology of the c e l l c y c l e . Cambridge University Press. Parsons , T.R. and R.J. LeBrasseur (1970). The a v a i l a b i l i t y of food to d i f -ferent trophic l e v e l s i n the marine food chain. In 'Marine Food * Chains' Ed. J.H. Steele. Edinburgh. Oliver and Boyd. 325-343. _and J.D. Fulton (1967). Some observations on the dependence of zooplankton grazing on the c e l l s i z e and concentration of phytoplankton blooms. J . Oceanogr. Soc. Japan. 23: 10-17. Pavlova, E. and L. Lanskaya (1969). Energy expenditures for movement i n some Black Sea d i n o f l a g e l l a t e s . 3rd International Protozoological Congress Leningrad. Progress i n Protozoology 3_: 182-183. Pavlovskaya, T.V. (1973). Influence of feeding conditions on the rate of food consumption and the time of generation i n c i l i a t e s . Zool. Zhurn. 52: 1451-1457. 185 Pearre, S. (1973). V e r t i c a l migration and feeding i n Sagitta elegans. Ecology 54: 300-314. Poulet, S.A. (1973). Grazing of Pseudocalanus minutus on n a t u r a l l y occurring p a r t i c u l a t e matter. Limnol. Oceanogr. 18_: 564-573. (1974). Seasonal grazing of Pseudocalanus minutus on p a r t i c l e s . Mar. B i o l . . ( i n press). Rapport, D.J., J . Berger and D.B.W. Reid (1972). Determination of food preference of Stentor coeruleus. B i o l . B u l l . 142: 103-109. Ricketts, T.R. (1971). P e r i o d i c i t y of endocytosis i n Tetrahymena  pyriformis. Protoplasma 73.: 387-396. (1973). The r e l a t i o n s h i p between endocytosis and dige s t i v e enzymes i n .Tetrahymena. J . C e l l B i o l . 59: 344 . Sheldon, R. W. and T.R. Parsons (1967). A p r a c t i c a l manual on the use of the Coulter Counter i n marine science. Coulter E l e c t r o n i c s Sales Co. Canada, Toronto, pp. 66. Steele, J.H. (1964). Some problems i n the study of marine resources. ICNAF Environ. Symp. Rome 1964, Contrib. No. C-4 pp. 11. Strathmann, R.R. (1971). The feeding behavior of planktotrophic echinoderm la r v a e : mechanisms, reg u l a t i o n and rates of suspension - feeding j . exp. mar.. B i o l . E c o l . 6_: 109-160. , T.L. Jahn and J.R.C. Fonseca (1972). Suspension feeding by marine invertebrate larvae: clearance of p a r t i c l e s by c i l i a t e d bands of a r o t i f e r , pluteus, and trochophore. B i o l . B u l l . 142: 505-519. Swift, M.C. (1974). Energetics of v e r t i c a l migration i n Chaoborus t r i v i t t a t u s larvae. Ph.D. t h e s i s , U n i v e r s i t y of B r i t i s h Columbia. 148 pp. Tappan, H. and A.R. Loeblich (1968). L o r i c a composition of modern and f o s s i l Tintinnidae ( c i l i a t e protozoa), systematics, geologic d i s t r i b u t i o n , and some new t e r t i a r y taxa. J . Paleontology 42: 1378-1394. Taylor, F.iJ.R. , D.J.Blackbourn and J . Blackbourn (1971). The red-water B c i l i a t e Mesodinium rubrum and i t s 'incomplete symbionts': a review including new u l t r a s t r u c t u r a l observations. J . Fis h . Res. Bd. Canada 28: 391-407. Theilacker, G.H. and M.F. McMaster (1971). Mass cul t u r e of the r o t i f e r Brachionus p l i c a t i l i s and i t s evaluation as a food f o r l a r v a l anchovies. Mar. B i o l . 10_:- 183-189. Throndsen, J . (1973). M o t i l i t y i n some marine nanoplankton f l a g e l l a t e s . Norw. J . Zool. 21: 193-200. 186 V i t i e l l o , P. (1964). Contribution a 1'etude des Tintinnides de l a baie d'alger. Pelagos I I : 5-41. Vlymen, W.J. (1970). Energy expenditure of swimming copepods. Limnol. Oceanogr. 15: 348-356. Williams, F.M. (1971). Dynamics of microbial populations, pp 198-268 i n 'Systems analysis and simulation i n ecology'. V o l . I Academic press Ed. B.C. Patten. (1972). Mathematics of microbial populations, with emphasis on open systems, pp 395-426 i n 'Growth by intussusception'. Trans. Connecticut Academy of Arts and Sciences 44 Ed. E.S. Deevey. Zaika, V. Ye. (1972). Microzooplankton of the Mediterranean and A t l a n t i c " Ocean o f f Northwestern A f r i c a . Oceanology 12: 408-414. and T.Yu. Averina (1969). Proportions of i n f u s o r i a i n the plankton of Sevastopol Bay, Black Sea. Oceanology 8: 843-845. _and N.A. Ostrovskaya (1972). Pattern of d i u r n a l v a r i a t i o n s i n microzooplankton abundance i n the surface layer of the Mediterranean sea. Oceanology 12} 725-729. Z e i t z s c h e l , B. (1969). 'Tintinnen}des westlichen arabischen Meeres, Ihre bedeutung al s indikatoren fur wasserkorper und g l i e d der nahrungskette' Sonderdruck aus "Meteor" Forschungsergebnisse Reihe 4: 47-101. Zenkevitch, L.A. (1963). Biology of the seas of the U.S.S.R. Press of Acad. S c i . U.S.S.R. Moscow. Appendix 1. Coulter Counter data.Tintinnopsis subacuta on natural p a r t i c l e s . N.T.C. method. 01 0. 1640E OS 0.1 2 70E 43 6. 0 05 13 7.0 0.3<>5BE 05 JV* 41 8F 05 0? 0,B3?3C 0.831 6E 0.3J16F 0, 63b OE O.6330E Oft Oh 06 06 06 06 Of. 0. I'fcOTE 05 0.2741E 05 _£i ial« 3?>3. 1624. _ 1 9 4 3 , 1 5 ,16 . 1 747. 2434. 0.5 15'IE 06 0,6)>7F 06 JJ-: SJ»!IE 06 03 0.5989F 0.KP65E 0.5B65F 0.6376<= 0.5705F o.6iri5r 06 U 6 36 06 06 06 06 04 4.000 8. 00 U 6.000 9. 000 2.000 4. 000 3. nop 0.6 >'. 6= OS 0.6T46E 06 3,. 6 I'M r 06 ? 6 5 « . 53?!. TOO. 6 514,0 51 T4. _3_9 09._ ? 2 4 ) . 8555. _5?43, 5106. 0.1261E 05 B2B?. 3900. 6946. _2X-_3._ 648?. 4364. -16.9JU 0,b393E 06 IF 06 _0.45-nP 06_ o.4 'i-»e"~ot> 0.471 IF 06 0,4_'7?E J6 0 . 4 > ? 7 F 06 0. 3526E 06 0 , ? > : . 5 F 06 0,25'. 5E 05~ 0.26130 06 _ 0 O i l 5? _06_ 0,2 )"'»£ 06 0.3174F 06 Q6_ 0. 3 377C 06 0 . 3 < 7 0 r 06 0.3379E 06 0.4742F 0.5414F _0.,f 414F_ 0 . 4 0 A 6 F 0.4966F 0.41 9 5r 06 06 06 06 06 06 2.000 8c 000 -J«oog_ 8. 000 8.000 1.1)00 0.4135E 0.3 64 7* _0.3f 47F 0. 0.1962T 0.3 5C6E 06 06 06 06 06 06 8,000 7. 000 _7.000_ 7.000 7.000 8.000 0.3506F 0.3200k" _0. 2 73 6 r 0.? 578F" 0.2 579F 0.2 57SF "0.2915F" 0.791. 5E _0.?5! 5|_ .1.3 517F 0.3 597E 0.35Q7F 06 06 06_ 0 6 06 06_ 06 06 06 7.000 5.000 5. 000 3. 000 B.OCO _R. 000 *7boo" 8. 00 0 8. 30 0 0.3 )>9E 06 0 .3H9* 06 . 0.3114E 06 06 6.000 06 8.000 06 7.000 0.3295E 0.3285F 0.2870E 0.3114F 06 0.3H4E 06 0.24K8F 06 2777. I 300. _N»M = MEANS 0.2. 970F 0.2370E 0.2 590F 06 6.000 06 7.00 0 06 6.000 06 8.000 06 8.000 06 6.D0Q 05 0.6200 0. 8500"" 0.6200 0.8 500 ' .000 0. 8500 1 . OOP . D6 0.4800 "0.4800. 0.7800 0.7800 1 .890 1.290 ) . 470 0. 8700 ] .490 i . 4 9 0 r 1.720 1 . 4 0 p 6. 300 0.2800 _J).7 800_ 0.5300 0.5300 4.110 1.72 0 l . * 9 0 1 .72 0 1.490" 1.72 0 1.490 4.110 5. 680 5. 680 4.840 " «. 840 2.710 1. 72 0 0.9200 O.o? 00 ] . 0<- 0 1 . 520 1 . 3P0_ "T.'dfrb 1 .470 1 . 06 0 1.470 1.170 2. 710 0.2000 0.7100 1.38 0 1.520 1.06 0 0. 64 00 0.6400 _0.6400 0.4 Vdb" 0.4300 0. 43 00_ 0739Q0 0.3900 0. 3900 1.470 1. 170 1. 380 0.4800 0.4800 0.7600 0.7600 0.7600 1.030 07 0.7700 0.7700 0.7700 0.7700 0.7700 0.81.00 O.A700 0 . « ? 0 0 0.S200 __0. 8 200_ ood l , ooo 0.8700 0,8700 0.8 700 0.8700 "6.8600 O. 8',00 0,8100 0.B100 0.8700 O.oinq_ 0. 8600 0. 8600 JO. n'-O0_ b . 6 tod 0.6600 _0.6600_ 0'. 6~h"6b 0.6600 0.6 600 0.8700 0.B7O0 0 . « 1 0 0 0. 81 00 0.9300 08 9. 000 9. 000 9. 000 9,000 10. 00 10.00 10.00 09 1 2.50 12.50 1 2.50 12.50 15.00 15.00 15„30 9.000 8. 000 8. 000 "a.ood" 8. 000 8. 000 0)0 '4 , 00 46.00 46.00 46. 00 46, 00 34,00 34 . 00 16. 50 18.00 1 « . 0 0 "18.30" 1 » . 3 0 18.00 8. 000 8. 000 8. 000 8.000 8. 000 8.000 1 8.00 18.00 18.00 18.i)0' 18.00 1 ° . 3 0 29* 00 42.00 '2.30 42,00" '2- 00 42, 00 8. 000 a. ooo 8, 000 8, 000 8. 000 " 8."000 8. 000 8. 00 0_ "ai ooo 8.000 8. 000 18.00 18.30 18.00 2 7"..3 0 27.00 77.00 77^00 27.00 27.00 "2 7. 00 27.00 27.00 42. 00 '2,30 00 42,00 42, 00 '2.00 42, 30 48, 00 _48. 00_ 9 0, 00 90. 00 90. no a. ooo 8. 000 _8.000_ 8.000 8. 000 8 . 000 01 _0 2_ 03 04 r>5 06 ~ D7 na 0* 010 5. « 7258.->5 484384, 4092 76. 6.43587 1 .2 7 614 1.5633? 0.814615 8,4615 3 0.2VISE C6 0.2590E 06 7.000 1 . 520" VT6?0" 0.9300 9.000 0.7315F 06 0.7715F 06 4.000 1.170 0.6100 . 0.6100 15.00 SISaOfV. CHRP FLAT IONS _ 39. OBSEPVATinNS 03, 04 05 06 OT OT" 27,00 27.00 27. 0O_ "27. 00 27.00 27.00 V 30 36.00 "•6. 00 36. OO 36. 00 34,00 10702.5 Jl 94.207. 01 1 .0000 0. ,'566 02 27.00 28.30 "09" 90, 00 90. 00 56,00 36.00 36. 00 90. 00 90. 00 29.00 ~onr 13818P. 1.90283 0. 3'"35 8 1.0000 21.1153 43.9742 S TM8F0* 1.7P807 0. 133599F-01 1. 21216 5.54348 19.9822 0.2662 0.4455 0 . 2 ° 6 ° _ -0."3~490 0.1283 -0.1713 -0.1602 -0.1253 0.9137 •0.0644 •0.2723 0.1134 •0.02.73 0.4453 •0.8025 1.0000 : -0.2855 1.0000 '71 0.5 801 1. 0000 0.0101 -0.0777 0.7968 CoWO" -0. 1523 0.1063 0. 3382 0.2434 0.7087 -0.4585 -0.3471 -0.0687 1.0000 -0.3625 1.0000 •0.4534 -0.634J 0.1286 0.2661 -0.3557 - 6 . 2 6 6 3 - 6 .1642 1.0000 •0.4291 0.0508 0.1595 -0.2749 0.43) 8 -0.2539 3.3630 1,0000 co Appendix 1. (Cont'd) ... >— 01—!_=1M5. • O . I R T ^ F - O J * n ? 0.«*00E . 0.35 O O F 05-05-1 FOamtCOEFF.M 5, »90 FPAOMCOEFe.)* 0.3^6 — • . < STO.^ R.CnFFF. =. 0.30A1E-02 STD.F«q. m . 0.2600E 05- 2 — • — RSQ = 0.1271 D I IR f l lN -MATS i lM ST».= 0.8675 AUTQCCfRELATtONCHE^ F. . Q.S??0 j 0. 17 00E 05-1 ] . * ' 8000. 1 2 I .1 I 1 I I 1 21. 1 11 • • 2 11 ? * -1000. 1 1 1 1 n~i 2 n 0.23QOE 06 0.3500E o.«70or 06 o;"fT6bT~o<S — 0 6 0.5900E 06 0. 8300E 06 i C 0.4400E 05- 1 > — -HI 3 - 7 8 6 . 3 • 3 . 1 9 6 6 E - 0 T * l» 0.35 0 0 E 05-1 FDATTHCOEFF. )=. ?. 8'3 EPRL'illCOEFF. I« 0 . 0 1 7 6 j < STD. F R R . CDVS T . a 5 0 4 7 . STr>.r»R. C O ^ F F . « 0.1170F-01 0.26 OOF 05-2 RSU » O . 0 7 0 9 — — • OURIlN-WATSnN ST*.« 0.9716 . ... AimCU'RELATION CDs**. « 0.5319 0.1700F 05-1 1 1 < j 8000. 1 1 1 1 1 > 2. 11 t" 1 11 11 1 • 1 21 1 2 i> 1 1 " 1 1 J 1 1 1 1 -1000. 0.2200E 06 0. 3300E • 0.4400E 06 06 0. 0.S600E 5500E 06 06 0. 7700E 06 00 00 Appendix 1. (Cont'd) 0.4400E 05- 1 > — DJ -DIM. » 7389. » n« 0.35 OOF 05-1 FRATTOICOEFF.). *,161 F C R T H i m F F F . ) . 0.0045 j • < STO.FOR.CONST." 5?91. STO.EKR.CDFFF.- » ^ , 2 S T n . P R P . ni « 1>57. O.PfOOF 05-2 1 R s i) = o.i9i>; 1 — ; 0'J<"UN-WAT?riN STA. - 0.6267 1 A"IOr.ri»"CLAT ION OI-i-F. • 0.6860 | j 0.17 00E 05- 1 i 8000. | . 1 * 1 j 2 1 • . 2 2 1 4 1 A 3 -1000. 1 2 2 . 2 • 1 1 2 \ 2.000 3.400 4. 800 6.200 7.600 9. 000 i co Appendix 1. (Cont'd) c \ ni > -4658. • 9333. * ns fl l « 0.1926F 05* -1418. • OR > f FRATIOtCOEFF. I» J, 576 FPRORCCOEFF. 1= 0.0614 FRATIOICOEFF.I- 1.118 FPRORICOFFF. 1" 0.'977 s. STO.FRR.CONST." 6497. STO.ERR.COEFF.. '4)33. S T n . F R R . 01 » 9873. STO.FRR.CONST." 0.1146E 05 STD.FPR. COrFF." 1341. STO.FRR. 01 * 0.1019F 05 RSO « 0.0 9 31 0UR!3IN-WATS0N S T A . 3 0.7023 AUTOCORRELATION C0=-F. " 0.6443 PSO • 0.0273 1 OURBIN-WATSON STA. » 0. 3225 AIITfirnPRFI A T T D N f.OF" F . = 0.5963 ni • 0.1037F 05* -1991. * 06 01 » 0.1 348E 054- -294c 9 * 09 \ • " FPATIOICOEFF.|. 5.1.31 FPROrt{COEF F.1" 0.3290 FRATIOI COEFF. |a 0.9746 FPRD8(C0EFF.»" 0.3313 1 STO.ERR.CONST.. 2373. STD.ERP.COFFF." 379.1 STn.EPR. 01 = >S99. STO.ERR.CONST." 6515. STO.ERR.COEFF." 293.7 STO.FRR. 01 * 0 .1 021E 05 PSQ » 0.1213 DURKIN-WATSON STA." .0.6177 AUTOCORRELATION C>3~F. = 0.6763 PSQ * 0.0257 DUPOIN-WATSON STA. • 0.3434 AUTnr.ORRFI A T ION f . n « F . a 0.5706 • 01 . -4104. + 0.1 395E 05» 07 01 • 0.1039E 05» -63 .C9 * 010 FRATIOICOEFF.l" 0.6194 F P R n B t r n F " . >. o . 44H FRATin(COEFF.|. 0.5902 FPROBIf.OFFF. !• 0.4531 STO.FRR.CONST." 0.I4.3F. 05 STO.FRR.COEFF. • 0.1 772F 05 STO.FRR. 01 " 0.10->5F 05 STO.FPP.CONST." 4397. STO.FRR.COEFF." 93.28 S T O . F R R . 01 " 0.10'6F 05 RSO • 0.0165 0UR81N-WATSJN STA.= 3.8449 AUTOCORRELATION COEc F. " 0. 5746 PSO • 0.3157 OUR BIN-WATSON STA," 0.8640 A t J T i m R R F I A T lf;\1 F n F C F . « 0. 55 95 . , -VO o Appendix 2. Coulter Counter Data. Tintinnopsis subacuta on natural particles. E.S.O. method. r,j 03 D4 05 06 07 DP 09 010 ?3">H. 0 . 4 ? 3 7 5 Of* 0 . 3 3 0 6 8 0 6 4 . 0 0 0 0. 5"00 0 . ] i s 4 0 E 0 1 0 . 1 2 7 0 F 0 5 1 7 1 5 . EAQ.O \ ) . 1 S ? 5 E 0 6 0.7117' 0 6 0 . 8 It o c . 0 6 3 , 1 6 W F 0 6 0 . 2 ? f 6 F 0 6 Q,-)77->= OH 0.7 5 7 6 E 0 6 0 . 5 C 7 1 T 0 6 0 . 5 8 6 5 8 O A 0.1477F 0 6 0 . 2 047<= 0 6 o. - ' r . ' .n 0 5 8 . 0 0 0 6 . 0 0 0 9 . 0 0 0 2 . 0 0 0 4 . 0 0 0 ] , 0 0 0 0. 8 5 3 0 0. 6 5 0 0 0 . P 5 0 0 1. 3 5 0 1. 4 5 0 ?.?io 0 . 4 8 0 0 ~ 0 . 4 S 0 0 " 0 . 7 8 0 0 0 . 7 8 0 0 1 . 3 9 0 1 . 2 9 0 O . " 2 0 0 J 3 . 7 7 0 0 0 . 7 7 0 0 0 . 7 7 0 0 0 . 7 7 0 0 0 . 7 7 0 0 0 . 8 1 0 0 o.apoo 9.000 9.000 9.000 9. 000 lo.oo IO.OO 10.00 12.50 12.50 1 2 . 5 0 1 2 . 5 0 1 5 . 0 0 1 5 . 0 0 1 5 . 0 0 ? 6 5 1 . 0 , 333 0 6 H 7 . 0 0,•')?''>" OS .J.3R5JE_05 3,6 >3?»..06. 0 . 4 < , ! 8 F . 0 5 3 i 6 •>>•"? 0 . 2 6 9 7 F 0 5 0 . 5 5 » 6 C 0 6 •1, .- '61 8 Q S O . C S 06 0 . 2 4 1 6 = 1 6 0 . 6 3 ? 3 " 0 5 _a . .5 .4 ! .4F . _0 .6_ 0 . 5 4 1 4 = 0 6 0 . 4 9 6 5 = 0 6 Q.A'Sr . 4 ' " O A 3 . 0 0 0 2 . 0 0 0 _?°oo.o._ " R . 'bod" 8 . 0 0 0 8 . OOP 1.0)0 0. e300 1.4"0 7 ? 6 1 ,4°0 1. 7 2 0 1 . 4 7 0 6 . 3 0 0 _0.2P.OO_ 0 . 2 8 0 0 0 . 5 3 0 0 0 . 5 3 0 0 0 , 8 6 0 0 o.proo 0 . ° 2 0 0 _ " b . ° ? o o l o 0 0 0 1 . 0 0 0 10,00 o . O O O 8. 000 ' » . o o b " 8, 000 8. 000 1 5 . 0 0 1 6 , 0 0 . I R o O O _ T f . o b 1 8 . 0 0 1 8 . 0 0 ? 0 4 7 , ?3! 3. J 6?4,_ 1 5 0 5 . J251,. 3 . 6 i n c 0 . - V 11 = 0 6 0 6 0 6 0 . 4 ? 1 3 c 0 6 0 . 4 7 1 O E • 0 6 3 i 4 7 J 3 = 0 6 O . M R I E 0 6 0.4.18 5': 0 6 . _0« v_ft7!L_o<L, 0"."T>-47c O f 0 . 3 - ! 6 2 = . 0 6 o . 3 q < - 7 c f )6 8 . 0 3 0 8 . 0 0 0 _7. pO_0_ 7 , 0 0 0 7 . 0 0 0 7 . 0 0 0 1 . 4 9 0 1 . 7 2 0 _1. 4 J * 0 _ r . 7 ? b 1. 4 9 0 1 . 7 2 0 4 . 1 1 0 4 , 1 !0 5. 6 8 0 0 . 1 7 0 0 0 o 8 7 0 0 Oo 8 7 0 0 8. 000 8. 000 8„ 000 5. 6 8 0 4 . 8 4 0 4 . P 4 Q 0 . 8 7 0 0 0 . 8 6 0 0 0 , 8 6 0 0 8 . 0 0 0 8 . 0 0 0 8 . 0 0 0 1. P., 0 0 1 8 . 0 0 H I , 0 0 _ '1 8 . 0 0 i».oo 1 8 . 0 0 7 4 3 4 . 2 6 6 8 . .715.0.. 6t . 1 i . " 1 > Q _ S 3 ? 8 . ? 5 1 1 , _ " f 5 5 5 . ' 5 2 4 3 . 51 0 6 . 0,4. '7 7F 06 9 . 4 ! ? 7 F . 0 6 0 . . i i i u os "o.'? »11E' 06" 0.1 6? IF 06 j j i - H j r o h 0 . 3 5 0 6 = 0 6 0 . 3 5 0 6 " C 6 0 , 3 is? 5 " 0 5 "o . i 577= o i " O o ! 4 3 5"= 0 6 0.1 0 7 IF. 0 6 8 . 0 0 0 7. 0 0 0 J . 0 0 0 5, 0 0 0 5. 0 0 0 lo 0 0 0  . 4 9 0 1. 72 0 . .1 . 02 0 " b . 9 i do 0 . 9 6 0 0 1 . 1 3 0 7 . 7 1 0 2 . 7 1 . 0 0 . 2 7 0 0 ~ 0 , " 2 0bb ' 0 . 7 1 0 0 0 . 6 4 0 0 0 . 8 1 0 0 0 , 8 1 0 0 _ 0 o K 7 0 0 0 . 8 7 0 0 " O.o? 0 0 0 . P 6 0 0 8. 0 0 0 8 . 0 0 0 a, 0 0 0 3'.""00'6" 8, 0 0 0 P . 0 0 0 18.00 1 9.00 1 P.00 "18.00" 1P.00 1.8 . 00 0 , ? 2 J ? = 0 60 . ? ? > 2 F Oi g.) r ' . 9 E _ 3 4 ' 0,3 I'o? 0 6 0 . 1 ) ' 6 F 0 6 0,317OC qt, 0 . 2 2 6 4 ' : 0 6 0 , ? ? 6 4 F . 0 6 0 . 1 4 O 7 E _ _ 0 6 _ 0 . 2 ' ' : 5.1 0 6 0 . 2 91. 5 F 0 6 0 . 3 5 1 7 F 0 6 8 . 0 0 0 8 , 0 0 0 4 . 0 0 0 _ " p","bod 8 . 0 0 0 6, 0 0 0 2 . 0 4 0 !.. 3 8 0 l o 0«0 1 . 4 7 0 1 . 1 7 0 1 . 0 6 0 0 . 6 4 0 0 0 . 6 4 0 0 _ 0 . 4 3 0 0 _ 0""."4'3bb 0 . 4 3 0 0 0 . 3 9 0 0 0 . 8 6 0 0 0 . 8 6 0 0 0 . 6 6 0 0 0 . 6 6 0 0 0 . 6 6 0 0 O 0 6 6 O O P . 0 0 0 8„ 0 0 0 _ P , 0 0 0 B. bob" 3 , 0 0 0 8 . 0 0 0 2 7 . 0 0 2 7 . 0 0 _ 2 7 . 0 0 _ 2 7 . 0 0 2 7 . 0 0 2 7 . 0 0 0 . 1 2 6 1 E 0 5 8 7 ? 2 . . 1 ? 17. 4 4 1 6 . 7 0 " 1 . 0 . 3 U 9 ? C 4 0 , 3 37 IF 0 6 0 „ ! 2 < 8 C 0 6 " O c 2 2 i VF'OST O . ^ ' l ' F 0 6 0,?VJSF. 0 6 0.?5°7: 0 6 0 . 3 5>'F. 0 6 0 . 1?'. 7" 0 6 _ 1.'2 7' :»5= ;"b6 ' 0 . 2 9 7 4 C 0 6 0 . 2 2 7 1 F 0 6 8 , 0 0 0 7. 0 0 0 3 ^ 0 0 0 "i' .Obo" 7 . 0 0 0 6 . 0 0 0 1 . 4 7 0 1 . 1 7 0 1. 1 5 0_ " U ' s T d 2 . 04 0 1 . 0 0 0 0 . 3 C 0 0 0 . 3 9 0 0 _ 0 . 4 8 0 0 b ' . 4 ~ 8 0 d ~ 0 . 4 8 0 0 0 . 7 6 0 0 Oo 6 6 0 0 0 , 6 6 0 0 _ 0 , 8 7 0 0 C . 3 7 0 0 ' 0 . 8 7 0 0 0 . 8 1 . 0 0 6 ' . " 2 . 4 ' •>4. 4 M . 0_ 2 " 7 7. C ! , 3 U -VF OS 0 . I l l 4 F 0 6 0 , ^ ) 3 8 F _ 0 "•_ " b , l 0 6 0 . 2 i " ' 1 ? 0 6 P , 3,16 3 M n 0 , 7 - 7 0 - 0 6 0 . 2 8 7 0 E 0 6 0 . r - . ; S 3 ? 01 0 ' . i ' 3 i ~ < R 0 6 •1 . 2 33 )!? 0 6 0 .7 1 r ' 6 E 0 6 8 , 0 0 0 8 , 0 0 0 2 . 0 0 0 6 . 00 0 7, 000 4„ 000 1 . 4 7 0 ! . 1 7 0 J . ) . 7 0 _ 1 . 0 0 0 2 . 0 4 0 1 . 2 7 0 0 . 7 f . 0 0 0 . 7 6 0 0 1 . 0 3 0 0.PI 00 0.8100 • O . i ? O 0 8 . 000 8 . 000 8 . OJ) 0 "'80" 000" P . 000 8,000 80 000 2 7 . 3 0 2 7 . 0 0 2 7 . 0 0 _ " 2 7 . 0 0 2 7 . 0 0 2 7 . 0 0 2 7 , 0 0 1 , 0 3 0 1 . 0 3 0 Oo 6 1 0 0 0 . 9 3 0 0 0 . ^ 3 0 0 0 . 6 1 0 0 8 . 0 0 0 p.. noo ~8Toe"o~ 8 . 0 0 0 15. 00 2 7 . 0 0 2 7 . 0 0 " 2 7 . - 0 0 " " 2 7 . 0 0 2 8 . 0 0 1 6 7 7 . 01 02 0) 0 . 1 " ) 1 3 C 0 6 0 . 1 2 6 7 J 0 6 1 , 0 0 0 0 . 9 7 0 0 • . f A N S S T O . O E V . COP.K F L A T I ON 5 . _ ni n_2 nj '67e~5".~5) o " 7 0 7 . " 8 4 " " " ' '"i."ooob 3 6 3 4 8 9 . . 3 2 S V I 6 . 0 . 5 8 1 7 1 . 0 0 0 0 7 o " 0 7 ? . 1 5 1 ?_9 I . 0 o 6 1 _ 8 _ 0 _ 0 . 9 7 7 7 1 . 0 0 0 0 0 . 6 7 0 0 0 . 5 ? O 0 1 5 . 0 0 2 8 . 0 0 44. P B S F R V A T I O N S 04 0 5 0 6 07 08 0 9 46.00 4A.00 46.00 * A . 0 0 3 4 . 0 0 3 4 . 0 0 3 « . 0 0 3 4 , 0 0 2 9 . 0 0 4 2 . 0 0 ' 2 . 0 0 " 4 2 , 0 0 4 7 , 0 0 4 7 - 0 0 4 2 . 0 0 4 7^. 0 0 _ 4 7 . 0 0 4 ? . 0 0 4 ? , 0 0 4 2 . 0 0 4 7 . 0 0 4 P . 0 0 " 4 8 , 0 0 " " 4 8 , 0 0 OO. 0 0 O 0 . 0 0 9 0 . 0 0 3 6 , 0 0 3 6 , 0 0 " 2 6 , 0 0 3 6 . 0 0 3 6 7 0 0 3 6 , 0 0 9 0 . 0 0 " 9 0 , 0 0 " " 9 0 , 0 0 3 6 . 0 0 »4, OO 3 6 , 00 9 0 . 00 - < ? ' 5 ; ' 0 ' 0 -9 0 . 0 0 7 9 , 0 0 010 0 4 5 . ' 3 t < 6 3 6 0 5 l . - l 1 8 1 0 6 _ 1 . 4 6 2 ? 6 _ 0 7 " 0 . 8 1 S 9 J 8 ' 0 3 8 . 6 1 2 6 3 n J 2 1 . 1 1 38 2 . 1 3 4 2 1 0 . 3 ' i 0 : i 4 1 . 7 0 " ' 5 7 0 . 4 4 7 2 0 . 2 2 3 7 - J 5 . _ 3 1 0 6 _ 0„ 1.0'.': 0 . 6 7 7 2 0 , 0 8 6 ] 0.1 3 3 5 0 . 7 4 f 7 0 . 1 5 1 7 0 . 0 54 1 1 . 0 0 0 0 0 . 4 5 6 0 0 . 0 5 1 3 " 0 . " 9 7 3 9 1 7 E - 0 1 . 1 0 1 2 0 . 0 1 0 4 -OTOE? 0 0 , 0 6 5 1 1 . S 4 3 4 2 - 3 . 1 6 0 3 - 0 . 1 31 ? - 0 . 1 3 4 5 - 0 . 4 7 3 6 5.5 7 7 9 8 - Q . : 5 4 3 - 0 . 4 6 5 2 - 0 . 3 3 7 1 0 . 0 9 0 2 1 . 0 0 0 0 0 . 2 2 1 4 1 . 0 0 0 0 0 . 3 0 7 4 0 . 2 1 3 1 - 0 . 7 4 8 0 - 0 . 0 9 3 2 0 . 2 3 9 4 - 0 . 3 ? 4 3 I.oortiT - 0 . 4 7 8 0 - 0 . 7 9 1 0 "ilO 4 9 . 7 4 0 ' * 2 1 . 3 1 4 4 ,1404 - 0 . 2 9 6 6 -0.2591. 0. C0e2 0.249! -0.2205 0.47' 3 1.0000 0.07)6 -6. 32! 2 1.0000 0.3?3O—1.0000 Appendix 2. (Cont'd) t 0. 44 0OE 05- 1 V. ni „ - ? « ! < ) . » 0 . 3 5 1 9 P - 0 ] * D? 0. 35 OOF 05-1 f FRATiniCOEFP. )=- 7?.23 J ir . iQ ' . r tcoEfF-.) . :).3o,in J < STO. F'.R. CONST. = 3?*3. STO. l ^ . ' . C O E F F . * 0.5 379E-O2 S T U . r i j B . n i * 7 l/ , n . 0. 26 00E 05-2 FSO = 0.3V65 Plie<M'l-WATSO') STA.* •.»,t>785 . ..AUTProsoF.tATlnw r i :=F , » 0.6812 j < • 0. 1700E 05-1 1 * • • J . 1 • 8000. 1 1 . * 1 11 1 7. • 1 1 11 . 21 1 1 2 . 1 2 1 7 111 2 1 -1000. I 12 11* 1! 1 ). 0.1000E 05 0.3900F 06 0.2 000F. 06 0. i 0.7700F 5800E 06 06 0.9600E 06 r 0. 44 00E 05- 1 v . 01 , - 4 4 6 1 . * 0 . 3 7 6 6 = - 0 1 » 03 0. 35 00E 05-1 FPSTIK COEFF. I- 25.95 FP808(C0E.rF.)> 0.0000 J < STO. EPF.. CONST.- l't"2. s To.Fc;.cor< : ' ' : .= o.7 394n -o» 0.2600E 05-2 < "%Q = 0.3 319 PU9 . ! \ I N - W A T S I 1 V STA." 0. 6566 AUTOCORRELATION COJc,= . =» 0.6670 j • 0.1700E 05-1 1 0 1 1 1 •-• 8000. 1 . 1 * I 1 1 * ! 1 1 1* 11 3 1 ? .1 1.!. 2 21 12 1 -1000. 1 1111 111, 1 1 1 10000. 0.3100E 06 0.1600E 06 0. 0.61 OOf 4600E 06 06 O.7600<= 06 Appendix 2. (Cont'd) r ; ——— 0.4400E 05- 1 L 01 = -395" 4 . * 1821. * 04 •' 0. 35 00E 05-1 FRATIOICOEFF.>» 13.50 _ cPRO'UC0E':f=. 1 = 3.30.34 j < 5T-!,t^P. CONST.-. 3363. S T O . r^ R . C P E ' e , - 5S1.9 s ' - n . F ' . s . o ' « w . 5 . 0.2600E 05-2 »5.7 = 0 . ? 0 - . ) 0 1 Otl^IM-WATSON ST4,» 3.6889 - 1 W l H I S f H T I l M r.i)3FF. - 0.6533 I 0. 1700E 05- 1 ). 1 ' 8090. i 1 1 . < 1 1 1 1 1 1 . . 3 1 7 2 1 4 2 3 3 -1000. 13 2 . 2 1 1 1 1 1-2 1 1. 000 2.600 4.200 5. 800 . 7,400 9.000 Appendix 2. (Cont'd) c \ 01 " -1143.. • 602S. * 05 01 » 0.1 545E 05» -1003. * 03 J r FR4TIC1 (COEFF. )* 2.212 0.1405 FRATIO(COEFF.)= F p R 0 8 f C n F p E . 1 = I. 108 0.'991 < STO.FRR.CONST." STU.ERR.COEFF.= S T O . F O R . ni 5509. 4053. )5^4. STO.ERR.CONST." STO.FRR. COEFF. = S T O . F R R . m = 3380. '5 3.0 16 96. RSO • 0.0500 OURRIN-WATSPN STA." 0.7275 AilTOrrioRPLATinN COS=F. = 0. 6350 PSQ » 0.0257 DUPBIN-WATSPN STA, = 0.8217 AUTOOOOOFI A T lOM m = =F. = O. <!R7« • \ 1 01 = 91/-7. • -1766. * n6 01 = 0.1 2 49= 05* -271.3 » 09 I FRATIOtCOEFF. ) = „ . FQRnntCQFFF.). 4.454 0,33°i FRAT TO(COEp F » )« FPR08ICHFFF.)= 1, 025 0.1185 i ( STO^FSR.CONST, = STO,E'.R.COEFF." S T O . F R R . 0) 1 « 6 2 . 33 3.9 3137, STD.ERR.CO^ST." STO.ERR.COEFF." S T O . F P U . ni " 53 44. 268.3 9735. 1 | 1 I -RSO - 0.0 165 OU" 3! N-WATSON STA. - 0.6275 JUTQCOPR^LAT ION Cn=FP, « 0.6759 PSO • 0.0238 OUR BIN-WATSON STA." 0.3467 AUTOrORRFI AT10M C.O"=F. = 0-5711 t j 1 ! i 1 l 01 « -1467. * 0.1009F 0^* 07 01 » 9°6 3. • -44. 26 * 01 0 1 ! FRAT!0(COfFF. )» ?PSC3 C.CQ«f_«J"_ 0.4148 0,5 2 03 FRATTOfCOEFF.)-FPOH8(COEFF.|" 0.3 449 0.3664 ; S^O.E- R . CON ST. = STO.ERR.COFFF.* STO.FRR. 01 0.1 257E 0"= 0.1 5 30 E 05 977?. ST0.ERP.-.CONST.-STO.ERR.COEFF.. STO.FOR. 01 3774. 69.91 77'5. i P.SO > .0.0102 DUCRIN-WATSON STA," 0. .5364 AUT Of 1 R P ELAT ION COF=F. , 0. 5797 PSQ • 0.0197 OUR 81N-WA TSO N STA," 0. 3572 AUTDCORPFI ATION CO=FF. = 0. 5645 ^ — . : . . ' , Appendix 3. Coulter. Counter data. Tintinnopsis subacuta on laboratory food. N.T.C. method. Dl PI pr P t P* Pf PI -* , I V ? £ . * O f H l f l W « p I U . K t n «. 0 0 0 a. O. nloa 9. 000 IS. So Cx. 0 0 0 . 1 4 4 9 ? 05 O.' .MIE 07 0.1 1 4 6 F 0 7 6.000 O.oiOO 0.6400 . 0 . 8 3 0 0 10.00 1 5.00 24,00 > 4262. . 0.3155E 06 0.7477E 0 6 4.000 0.5500 0.7900 0.8100 9.000 25.00 7 4 . 00 3458. 0 , ! ) i ) F 0 7 0 . 1 9 ) 9 F 07 3.000 1 . 040 1 . 390 0.8000 10.00 2 5 . 5 0 30.00 17.00 0,1770? 07 0.' 507F 0 7 2.000 0 . 8 2 0 0 1 . 3 9 0 0.9600 10.00 25.50 3 0 . 00 970. 0 O.H03E 06 0.1231E 06 6.000 2. 8?0 2.580 0.6800 8 .000 74.00 36.00 > O51..0 0.3?'7F 04 i l . ' U l f 06 1.000 1 .350 0.8200 0.7200 8 .000 2 4 . 00 36.00 1298. 0, 6341F. 06 O.7 340F 06 5.000 1.31 0 1.450 0.7200 8 . 000 2 4 . 00 36. 00 11. .10 0 , ? 3 ' 3 E 06 0.;-513F 06 3.000 0, 5400 2. 230 0.3000 8 . OOO 25.00 74 . 0 0 > 5 1 1 . 3.41>3F 06 Q„3?47F 06 6c000 0.8400 1.850 0.7500 9. 000 25.30 2 4 , 00 N A M E MFAN5 s r n . n r v . • CORP FLAT IONS 10. 08SFRVAT IONS 01 02 D 3 04 05 06 D 7 0 8 D9 0 1 0 PI 4?7">.70 5402.62 1 .0000 02 901075. 7246?6. 0 . 4 6 3 3 1.0000 01 1245.2). 616477. 0 . 4 2 7 4 0 . 9 9 4 6 1.0000 04 4,00000 l ,763f* 0 , 3 » 3 5 -0,1827 - 0 L 1 9 4 5 JiOOOO I 0 5 1 . 11 300 0. 663425 -0.2.166 -0. 3278 -0. 2 75 6 0.2925 1.0000 — — P 6 1.35400 0.7081 7 8 -0.6680 - 0 . 4 5 5 5 -0.4345 0.742 8 0.4699 1.0000 r>7 0. 7 ) « J J J 0. 9 57I3 5F -01 0.2967 0.7330 0 . 6 6 9 3 -0.251 3 -0.6129 -0.4601 1.0000 03 8.80000 0.9!89"<5 0.4425 0.8327 0.805 7 -0.1371 -0.3488 -0.4777 0.8031 1.0000 P-* 22. 9500 4.J4.967 -0.9409 -0.3778 -0.35)1 -0.3345 0.0695 0.5931 -0 .3391 - 0 . 3 0 7 6 1.0000 P 1 3 , 1 . 6 0 0 J 8 , 8 8 * 4 3 0 , 7 7 7 6 0.2959 0.3276 -0 .1 276 0.341 8 - 0 . 2 8 9 9 -0.0834 - 0 . 1 1 9 8 - 0 . 4 0 9 6 1 . 0 0 0 0 * 5. •SIMP.r.G* 0. 14 5 0 E 05-1 1 v 1 > 0 1 « 1161. f 0 . 3 4 5 4 E - 0 7 * 0 7 1 1 0.1160F 05-* FR AT IO( COEFF. I * 2 . 1 8 6 1 1 < FPR09ICnEFF.I» 0.1754 1 STO.FDR.CON ST.• 2f47. 1 S T O .FRR.C0FFF.= 0 .2336E-02 8 7 0 0 . S T o . ^ o . 01 • 5,173. 1 < PS ) = 0.2146 1 nn«3IN-WATSCN ST\,= 0.7702 1 AIJT'XOOREIATION r,o;=F. » 0 . 5 1 6 5 1 • 5 8 0 0 . 1 1 • • 1 1 1 1 • • 1 2 9 0 0 . - • 1 1 > . . ! 1 1 1 -0 .0 1 1 X 0 . 9 0 0 0 E 0 5 0 . B 3 0 0 E 0 6 0 . 1 5 7 0 E 0 7 0 . 4 6 0 O E 0 6 0 . 1 2 0 0 E 0 7 0 . 1 9 4 0 E 0 7 Appendix 3. (Cont'd) r 01 » 1539. • 0.3315F-02* 03 0.1A5OE 0. 1160E 05-1 1 1 1 05-1 1 < > FRATI0(CnEFF.)« 1.788 FPRO3(C0EFF.)• 0.2165 1 1 1 STO.FRR.CONST.' 36 2 0. STD.EPR.COEFF,= 0.2479E-02 STO.FRR. 01 » 1131. 8700. 1 1 < °S0 = 0,1827 OUR TfN-W ATSON STA.* 0.7043 MfTienPRCiATinN r.n^F, • 0,54^ 8 1 1 1 • 5800. 1 ! • • 2900. 1 I i I 1 1 I > 1 1 1 ' -0.0 l I 1 0.1200E 06 0.8400F 0.4800E 06 i 06 O.J200F 0.1560F. 07 07 0.1920E 07 -*25. 1175 . 0.1450E 05-0.1160E 05 FRATIOICnEFF.J. FPRORICOEFF.)» \. 3 71 0.2739 STP.ERR.CONST. « 4336. STD.ERR.COEF F.= 1000. sTn.FRR. oi » r > ? 9 - > . 8700. PSO. * 0,1 471 0UO<3IM-WATSPN STA.= 0.6175 _A I IT OC npRFt AT ION CQ-.rC. * 0.4690 5 800. 2900. -0.0 > I — 1.000 1 — I — 2.000 I 3.000 5. 000 A , ono 6.000 Appendix 3. (Cont'd) _ai_ 632,6. » -184? « 05 FRATIOtCOEFF.I- 0.4330 FPRQBICPEFF.I. 0.5345 STD.ERR.CONST.' 3586. STO.ERR.COEFF.- 2304. STP.FRR. 01 . 3591. PSO " 0.0513 OURBIN-WATSrN STA. • 0. 7*9* -A'JI.OCQR5ELAT10N CDEFF. * 0.4562 _D_1_ -0_1B.?F 0?» 26Q?. _____ FRATIOJCOEFF.) = FPRUMCnEFF.l . STO.FRR.CONST.-STD.ERR.COEFF.= S T n . F R R . 01 1.943 • 0.1995. 0.) 649E 05 1864. 51.39. RSQ « 0.1953 DUR8IN-WATS0N STA.» 0.6437 _AUT.QCOR.RELAT 1PN CQE ; F. » 0.4814_ _____ 0.111BF OS* -5096. _Q6_ FRATIOICOEFF.)• _FP.IO a Lcn_r_F_j__. 6. 447 O-T'38 STO.ERR.CONST.' 3037. STO.ERR.COEFF. a 2707. STO.FRR. 01 ' 4 7 6 4 . RSQ » 0.4463 OIJRBIN-WATSON STA. ' 1.804 AUTOCORRELATION COEFF. - 0.1124E-01 0.3296F 05> -1356. FRATIOICOEFF. I» _E£.RJJ_.LCi3EE.F. l» STO.ERR.CONST.' STO.ERR,COEFF.' STO.FRR. 01 61 .77 J..3001 3702. 159.8 1?TO. PSO ' 0.9853 OUR 8IN-WATS0N STA. » 1.013 AUTOCORRELATION COE-F. ' 0.4299 _ D J— . - o . i ' a a e os» o . ? n v 05« 07 FRATIOICOEFF.|. ..EERQ31.C0EFi_l__ 1. 494 STO.FRR.CONST.' 0.1650E 05 STO.ERR.COEFF.' 0.7P59E 05 STO.FRR. 01 . *jif>n. RSQ ' 0.1 5?4 DUR9IN-WATS0N S T A . ' 1.023 AUTOCORRELATION COEFF. ' 0.344? _Q__ - lQt - • 16 9. 8 010 FPATIOICOEFF.)> 0.1680 FPR0B1COFFF.I. O . 4 1 9 STO.FRR.CONS T.» 6755. STO.ERR.COEFF.' 206.5 STO.fRR. 01 ' 5505. RSQ » 0.0771 0UR8IN-HATS0N STA. » 0.6704 AUTOCORRELAT ION COEFF. . 0.5899 Appendix 4. Coulter Counter data. Tintinnopsis subacuta on laboratory food. E.S.O. method. PI p. *" 0<L_ 1?* ? 6'.T44«SE C5 C" 4682. 0, 3580. 0, 927.0 0. 1020. 0 167.0 0,  L 0 6 • 1625. 252.0 '140.0 420.0 367.0 569.0 3822. NAME MEANS P t <J >7le <n 8295E 06 2418E 06 3148E 06 2625E 06 ,1246fc 06 , 1085E_05_ , 752 aE 05 .3261E 06 lliVOL' 06 .1424E" 06 , 196 IE 06 73 15P C5 .4767E 05 •4323F 06 STO, Pi a.lotot 0.6874F 0. 1410E 0.4751E C.17S8E 0.1164E _0_.1213E_ 0.79 3 4E 0.3140E 0.2'->2 J U " " C . 1524E 0.1774E 0. 2566F " 0.37 74E 0.3247E OEV. P4-07 ».ooo 01 3671.32 02 29 3013. 03" 267005. 04 3.26666 05 1.57400 06 1. 74466 07 0.771333 OS 9.66666 0 9 21.3999" D10 36.3999 6190.55 319468. 2 78 59 9.' 1.83095 0.857535 1.46703 C. 834835E-01 _ 2 .2886') 5. 19684 10.9270 C6 6.000 06 4.000 06 3.000 C6 2.000 06 6.000 05 1.000 05 1.000 06 5.000 C6 3.000 C6~ 2. CCO 06 2.000 C5 1.000 05 """3.000 06 6.000 CORHELATICNS n i 0 2 1.00CO C.56CE 1.0CC0_ 6.9243 0.9727 0.43C9 0.5179 0.3334 -0.4144 •0.3984 -0.3174" 0.50C1 0.5127 •0.0662 0.08^6 ;6.3670 -0.3490 0.0313 0.C272 PS o. Woo 1.000 0.61 CO 2.860 0.54 00 . 2.930 3.110 1.650 1.150 _ 2 . 2 3 ( J _ 1.4 50 0.8700 1.620 ""' 1. 7 10 " 0.8400 03 1.00C0 0.4723 •0.2674 -0. 24 55 " 0.44 31 0. 16e0_ -0.2761 0.C6 84 P 7 PS l.oo Pt IS. ft fZ. oo 0.6400 0.7900 1.3S0 1.390 2. 580 1.600 0.8800 0.8100 0. 8000 0.9600 0.6800 0.6800 10.00 9.000 10. 00 10.00 8.000 B. 000 15. 00 25.00 25.50 15. 50 24. 00 24. 00 24.00 24.00 30.00 30. 00 36.00 36.00 0.82 00 1.450 _5.390_ 4.5 50 O.5SC0 0.5200 2. 2 3U 1.850 15. 04 05 0.7200 0.7200 _ 0. 6(100 "0.68 00" 0.7700 _0.7800 O.BOOO" : 0.7500 _CBSERVATIONS 06' 8.0 00 15. CO 15.00 9.0 00 " 9. 000 9.000_ ~0.0 00" 8.000 24.00 28.00 2(1. 00 " 15.50 " 15. 50 15. 50_ 25.00 25. 00 36. 00 37.00 37. 00 "52.00" 52.00 52. C0_ 24 .00 24.00 07 08 09 D10 1.0000 ^0.1709 l.0000_ -676008 0."3424 0.O863 -0.5902 0. 1591 -C.0393 0^2 207 0^4404 -0.4484 0.0123 1.0000 -0.5473 1.0000 0. 38 15 - 0 . 11 34 1^0000_ 6.3133 -0.51C9 0.3363 0.0455 -0.2528 0.0228 1.C000 -0.51 12 t.0000 01 •1784. • 0. 18626-01*02 FRATIGtCOEFF. )» FPR08IC0EFF.)= STO. ERR-CONST." STO.ERR.CGEFF." STC.ERR. 01 PSQ ""= 156.3 0.0000 _ 633.7 C.1489E-02 1780. "6.9232 OURRIN-WATSON STA." 1.602 AUTOCOKREl AT ION COEFF. => 0.9B13E-01 C.2200E 05-0.1700E 05 0.1200E 05 7 0 0 0 . 2000. 11 1. 1 11 .12 -3000. 1 -0.2000E 05 0.4600E 06 0.9400E 06 0.2200E 06 0.7000E 06 0.1180E 07 Append!* 4. (Cont'd) -1814. • 0.2054E-01* 03 FRATIOICOEFF.)-FPROB(COEFF.)-STO.ERR.CCNST.»' STO.ERR.COEFF." STC.ER8._01 __ a so -OUR 8 IN— Vi A T SON S AUVGCOKRCLAFION 76.28 o.cooo _ _ 891.7" 0.2352E-02 ? 4 5 l . 0.8544 IA.= 1.580 COUFF. - 0.1679 0.2200E 05-0.1700E 05 0.1200E 05 7000. 2CC0. 1 11. 1 2 - 1 1 1 >. -3000. -072O'B'OE 05 0. 4000E 06 0. 82 00E 0"6 0.1900E 06 0.61C0E 06 0. 1030E 07 Appendix 4. (Cont'd) 01 -1088. 1*57. * 0* 01 5403. • -179.1 • 08 FR AT IOICOEFF. )• FPRCBICOEFF.)-ST0.ERR.C0NST.»~ StO.FRR.COEFF.-STD.ERR. 01 HSO =• C. 1657 0UR8 IN-KATSON STA.» 0.6110 AUTCCORRELATI CN CCEFF. » 0.3485 2.965 0.1057_ 3143. 846.2 5797. FRATIOICOEFF. >-_ FPROB(COEFF.)__ STO.ERR.CONST.-STC.ERR.COEFF.« STO.ERR. 01 0.5726E-01 jp.80OO 7423. 740.5 6410. RSO = 0.0044 DURBIN-WATSCN STA.- 0.3226 AUTOCORRELATION COEFF. - 0.5385 01 » 7460. • -2407. * 0 5 FRATIOICOEFF. FPROB(COEFF. STO.ERR.CONST STO.ERR.COEFF _STD.F_RR._01  RSO DURB IN-WATSON AUTCCORRELATI 1.626 _0.2229_ '3358. 1888. 6057. » 0.1112 STA.- 0.6274 CN CCEFF. - 0.3980 01 —0.2493E 05* 0.3708E 05* 07 FRAT IOC COEFF. )• _ FPRCBICOEFF.)-STD.ERR.CONST.-STO.ERR.COEFF.-STD.ERR. 01 4.335 0.0554 0. 1381E 05 0.1781E 05 5563. ..01 - 0.1 LQ.3 _Jl?J____37__2_ FRATIOICOEFF. » _ FPROB (COEFF.j_ STD.ERR.C ONST. STC.ERR.CCEFF. STD.ERR. 01 • 09 RSO « C. 2501 DUR8 IN—WATSON STA.<= 0.8300 AUTOCORRELATION CCEFF. - 0. 3139 RSQ - 0.1347 OURBIN-WATSON STA.- 0.4674 AUTOCORRELATION COEFF. « 0.5060 01 6604. FRAT IOICOEFF. FPROB!COEFF. STO.ERR.CONST STC.ERR.COEFF STO.ERR. 01 RSQ 0UR8 IN-NATSCN A UTCC ORREI ATI -1681 . 2.452 0.1383_ 2413. 1073. 5893. » 06 = C.1587 STA.- 0.5371 CN CCEFF. - 0 . 4 5 4 7 01 3026. 17.73 » 010 FRAT 10 I COEFF. )» _FPR0B(C0EFF. }m_ STO.ERR ".CON S T . -STO.ERR.COEFF.• STD.ERR. 01 -0.1275E-01 _0-_8770 5952. 157.1 6421 . RSQ « O.C010 0UR8 IN-ViATSON STA.- 0.3012 _A_.LfiCORJlt LATI UN COEFF. - 0.5553 ts) O O Appendix 5. Coulter Counter data. Stenosomella ventricosa on laboratory food. N.T.C. method. Dl _1255._ 628.0 4114. 2797. 5378. 1022. 2829. 590 4." 959.0 464.0 1369. 2563. 5872. NAME MEANS Dl 2704.15 D2 649792. 03 723092. 04 4 . 0 0 0 0 0 05 1.4 744 1 06 1.96538 07 0.871538 OB 1 2 . 0 0 0 0 09 25.8461 U10 35.6973 02 0.5121E 06 0.5121E 06 0.6078E 06 0.6078E 06 0.6 12 IE 06 0.6121E 06 0.6121E 0.9336E 0.9673E 06 0.54C5E 06 0.61576 06 0.6698E 06 0.6441E 06 STO 06 06 0 3 0.7030E 0. 7030E 0.8023E 0.8023E 0.7562E 0.7562E 0.7562E ""C.9644E 0.8088E 0.5505E 0.6397E 0.6213E 0.6012E ^DEV. 04 3.000 4.000 3.000 3.000 4.000 5.000 4.000_ ~6. 000 3.000 2.000 7.000 5.000 3.000 CURRELA TI ON S 05 1.930 1.860 1.930 1.860 1.9 30 1.4 70 _ 1 . 8 60_ 1.1 50 0.7500 _ 1.2 00 1.3 CO ' 0.9400 0.9500 0 6 3 . 2 80 ' 3 . 2 80 1.410 1.410 1.140 1. 140 1.140 1.130 1.940 __5.850 2.940" 0.6700 0.2200 07 0.9600 0.9600 0.94 00 0.9400 0.9400 0.9400 _0.9400_ 0.8200 0.7300 _0-8900 0.8800" 0. 8300 0.5600 Dl D2 2C11.26 1.0000 141593. 0.2937 1. 0000 109496. 0. 3693 0. 5976 1.4 1421 C.1C47 0. 2358 0.441654 ' 0.0100 - 0 . 6159 1.51603 -C.6660 -0 . 36 52 0. 115319 -C.3878 - 0 . 5020 5.33853 0.2)67 ' 0. 8 I 14 1.5191 1 -C.2449 -0 . 1357 6 .7 74 75 -0.1829 - 0 . 0018 C3 1 .0000 C.2150 "0.2188 " -0.4187 0. 18 29 0.09 56 ' -0.5765 0. 57 14 04 J . 0000 -0.1508 -0.2748 0.0712 0".05 52 0.0388 0.4784 13. D5 OBSERVATIONS D6 0 7 1.0000 0.0716 1.0000 _0 i7910_ 0.33 75 1.0000 •0.0956 -"0.1866 -0.8060 1.0000 -0.4025 0.6684 -0.1359 0.1850 0 . 5 3 69 - 0.18 0 0 0. 6 7 72 - 0 . 48 62 1 . 0 0 0 0 - 0 . 5 3 1 3 1 . 0 0 0 0 01 -6.506 » 0.4172E-02* 02 5900. 4800. 1 FRATIOICOEFF. )« FPROSICOEFF. »« STO.ERR.CONST . -STO.ERR.COEFF.» STO.ERR. 01 _» RSQ = DURHIN-KATSON STA." AUTOCORRELATION COEFF 1.038 0.3318 2718. ~ " 0 . 4 0 9 4 £ - 0 2 7008. 0.0862 3 7 0 0 . 1.892 =• - 0 . 6 9 7 0 E - 0 1 2600. 08 09 o t o 8.000 25 . 0 0 39.00 8.000 2 5 . 00 39.00 J 8.000 25 . 0 0 39.00 8.000 25 . 0 0 39.00 8.000 25 . 0 0 39. 00 8.000 25 . 0 0 39.00 8.000 25. 00 39.00 ( 20.00 26.00 36. CO 22. 00 25.00 • 36.00 13.00 30. 00 22. 00 10.00 28.00 42.00 17. 00 26. 00 35.00 18.00 26. 00 20.00 D8 D9 010 15C0. 4 0 0 . 0 - 1 0 . 4 7 0 0 E 06 0 . 67 0 0 E 0 6 0 . 8 7 0 0 E 0 6 0 . 9 7 0 0 E 06 0 . 7 7 0 0 E 06 0 . 9 7 0 0 E 06 i Appendix 5. (Cont ld) ! j 5 9 0 0 . 1 ^ i l L 0 1 * - 2 2 3 5 . • 0 . 6 7 8 * 6 - 0 2 * 0 3 • 8 0 0 . j F R A T I O I C O E F F . ) - 1 . 7 3 7 ; F P P . O B I C O E F F . ) - 0 . 2 1 2 5 ! S T D • E R R . C O N S T . - " 3 7 8 6 . ~ — — " < i < ' S T O . E R R . C O E F F . - 0 . 5 1 4 7 E - 0 2 1 ' S T O . E R R . 0 1 = 1<552. H^D —> f\ 1 i t / 1 1 • 3 7 0 0 . J W U a I JO't D U R B I N - W A T S O N S T A . = 1 . 7 6 1 A U T O C O R R E L A T I O N r . O F F F - - - 0 . 9 3 5 2 E - 0 1 i • . i I 1 2 6 0 0 . 1 . : i 1 1 5 0 0 . • 1 > 1 i — 1 1 1 4 0 0 . 0 - — j - — 1 i - 0 . ! >200E 0 6 0 . 7 0 0 0 E 0 6 0 . 8 8 0 0 E ~ 0 6 ~ 0 . 6 1 0 0 E 0 6 0 . 7 9 0 0 E 0 6 0 . 9 7 0 0 E 0 6 5 9 0 0 . - 1 0 1 2 1 0 8 . • 1 4 9 . 0 * 04 4 8 0 0 . F R A T I O I C O E F F . ) F P R 0 8 I C 0 E F F . ) S T O . E R R . C O N S T . S T O . E R R . C O E F F . _ S T D . E R R • 0 1 RSQ OURB I N - W A T S O N S A U T O C O R R E L A T I ON 0.1220 0. 7298 ' 1801. 426.4 2C89. 0.0110 T A . » 1.852 COEFF. -_-0.58l5E-01 3 7 0 0 . 2 6 0 0 . 1 5 0 0 . 400.0 - 1 1 2.000 3.000 4.000 6.000 5.000 7.000 O IO Appendix 5. (Cont'd) 0 1 • ,637. • 4 5 . T 1 « 0 5 F R A T I O I C O E F F . ) - 0 . 1 1 0 8 E - 0 2 F P R O B I C O E F F . ) - 0 . 9 2 4 2 S T O . E R R . C O N S T . - 2 1 0 7 . S T O . E R R . C O E F F . - 1 3 7 3 . SVO.ERR . 0 1 2 101_-R S Q » 0 . 0 0 0 1 D U R B I N - W A T S O N S T A . « 1 . 8 6 0 A U T O C O R R E L A T I O N _ C O E F F . • - 0 . 5 T 1 2 E - 0 1 0 1 - 1 6 3 4 . • 6 9 . 1 9 » 0 8 F R A T I O I C O E F F . ) - 0 . 6 5 3 1 _ F P R O B < C O E F F . 1 « 0 . 4 4 1 0 S T O . E R R . C O N S T . - 1 4 4 0 . S T O . E R R . C O E F F . - 1 1 0 . 4 S T D . E R R _ C 1 ___ 2 0 4 1 . ; RSO » 0 " . 0 5 6 0 O U R B I N - W A T S O N S T A . - 1 . 8 6 B A U T O C O R R E L A T I C _ _ C O E F F . » - 0 . 2 2 8 3 E - 0 1 0 1 - 4 4 4 1 . • - 8 8 3 . 6 * 0 6 F R A T I O I C O E F F . I - 8 . 7 6 9 _ F P R O B ( C O E F F . ) • 0 . 0 1 2 6 S T O . E R R . C O N S T . - 7 2 9 . 9 S T C . E R R . C C E F F . - 2 9 8 . 4 S T O . E R R . D l _ » 1 5 6 7 . RSQ - 0 . 4 4 3 6 O U R B I N - W A T S O N STA.<= 3 . 0 0 5 A U T O C O R R E L A T I O N C O E F F . « „ - 0 . 5 5 3 2 0 1 - 0 . 1 1 0 9 E 0 5 » - 3 2 4 . 3 « 0 9 F R A T I O I C O E F F . ) - 0 . 7 0 1 9 F P R O B I C O E F F . ) - 0 . 4 2 4 4 lifb .ERRVtoNSf•« 0.-602. os S T O . E R R . C O E F F . - 3 8 7 . 0 _d___?R_»__01 - 2 0 3 7 . ' RSO « 0 . 0 6 0 0 D U R B I N - W A T S O N S T A . » 2 . 0 8 4 A U T O C O R R E L A T I O N _ C O E F F . - _ 0 _ _ J _ _ f 9 _ 0 1 8574. • - 6 7 3 5 . * 0 7 F R A T I O I C O E F F . ) -F P R O B I C O E F F . 1 -S T D . E R R . C O N S T . - " S T O . E R R . C O E F F . * S T D . E R R . 0 1 » 1 . 9 4 8 0 . 1 8 S 1 4 2 4 0 . " 4 8 2 6 . _ l _ 9 3 6 . _ . 1 5 0 4 RSO • 0 . D U R B I N - W A T S O N S T A . » 1 . 9 9 3 A U T O C O R R E L A T I O N C O E F F . - - 0 . 1 9 4 4 E - 0 1 D l - 4 6 4 3 . • - 5 4 . 3 1 * 0 1 0 F R A T I O I C O E F F . ) - 0 . 3 8 0 9 F P R O B I C O E F F . ) • 0 . 5 5 5 7 S T D . E R R . C O N S T . - 3 1 9 3 . S T O . E R R . C O E F F . - 8 8 . 0 0 S T D . E R R . 0 1 - 2 0 6 5 . RSO - 0 . 0 3 3 5 O U R B I N - W A T S O N S T A . « 1 . 8 8 7 A U T O C O R R E L A T I O N C O E F F . - - 0 . 1 7 8 8 E - 0 1 O co Appendix 6 . Coulter Counter data. Stenosomella ventricosa on laboratory food. E.S.O. method. F 5 U I 3 I CflFFF. ) = o . i n ' 9 J j.. - . - • ^ C . a K O F F F , 0,677' | .> i < .r <•-<.L.;'^ i . » 5 J 0 7 . ST0. c".P..C r!EFF J= D.O .T7F-02 . 9400. 1 1 . ^5) . 0 . J 1 2 1 ~ — — — _ _ _ r i ' . I R I I V-HiTSn*) STA," 1.1 OA 1 . * i ; r i C f , o o r i A T inf.- c.i«t. - o . ^ ^ - o i i 6400. 1 1 1 1 1 3400. I 1 . 1 -1 . 1 < l > 1 — — 1 — 1 1 1 1 400.0 1 1 1 1. 1 - 1 0.1600E 06 0.30C0F 06 0.4400C 0. 2300F 06 O.27O0F 06 06 0.5100F 06 O 4>-Appendix 6. (Cont'd) i —• — — :— \ 0. 1540E 05- 1 > oi - -09 1. ••-0.1S72E-0'* 03 0. 1240F 05-1 FRATIOfCOEFF.)- 0.^2-1 F-01 ...0.3 063 ! < .STO.FP R.TONS T . -STn.CRR.CCEF- F. = 2371. 0.31775-02 37 5 7. 9400. "SO = 0.3037 nilRBlM-WATSO"! STA.- 1.24? AUT„0$S5l4T. rN COS* e _ « _0JL2399F-0'. 6400. 1 1 1 1 > 3400. 1 1 1 < ! 1 1 1 1 1 1 '00.0 - 1 . 11 1 -1 / 0.1100E 06 0. l ° 0 0 E 0.2700F 06 06 0.3 5O0E 0.4 3 00E 06 06 0. 5100E 06 0.1S40E 05- 1 > > 01 = > M 1 . * - '7 .1 1 * 04 0.1240F 05-) FRATIOICOFFF.)-_ J J C n _ L _ n r c ; c , 1 * 0.2 3.70F-O2 0.-)\ M j < ST;i. F c " . CONS T. = STO.F'^.COEF n , . S T n . t ' c u . n i . 2734. ST 3. 8 17 6 ) , 9400. K i ' l = 0.0002 Dii*3iv-KA7SP ,I S M , » 1.133 A'JT0C._«.?...AT.I.yi.-C31 s e. » 0 . 4 ? l i r - o i j 6400. 1 ' 1 1 1 '- : 1 1 3400. • - » 1 ) • < 1 1 1 3 400.0 - 1 1 J 1 I 2.000 3.000 4. 000 5.000 6.000 7. 1 000 Appendix 6 . (Cont'd) * -V2.1 0.4 3 96E-01 0,3113 FPATIPlCOE^F.)" STO, ERR. CONST. - 2787. STO.EKP. COEFF." 1412. STO.tfP. O: 2 3757, _Q5_ J97JV PSQ " 0.0033 OUP 3 IN-WATSON STA." 1.215 ..AUT.OCOB?_p.LAT ION CO£ cP, " 0.36 33E-01 FPAT IO 'COEFF . ) . FPROn iCOEFF. ) . 0.5 275 0,4e54 STD.-PR.CPMST.- 22 56. STO.FP.P.COTFF. • 159.3 STO.FRR. 0) * 3605. PSQ " 0.0363 OURBIN-WATSON STA.= 1 . 1 9 3 AltTOCQPRFLAT ION CO'£ = F . * 0. 66 9 1 F - Q 1 6 3 4 " -l>5?i * 06 F P A T I O ( C O S " . I " FPP.03 <. C O r F C . ) = S T O ' . F P P . C O F S T , = S T D . p o o . C O E F F . " S T O . F O R . 01 4. 5 43 1353. 557.2 3' 06, RSO » 0,3192 011P31N-WATSON STA. " 1.739 _A_UTOC'IRPELAT ION COE=F. " -0.1972 _IU_ 0 . 2 3 6 4 ^ 0 * « ~ 0 . 7 7 9 3 F 0 5 « O f F O A T I O I C O E F F . I " 11.67 _ FTP . U D X C G F r F . I » 0 . 0 0 4 ? S T O . E R R . C O N S T . . • 5 ° ' , ' . S T O . F R R . C O E F F . . 471.2. S T O . E R R . 01 " • ' 7 3 0 . PS 0 = 0 . 4 5 4 4 0IJ°9IN-XATSON STA. = 1.063 -AUTOCORRELATION C'i;=F. " 0 , 3696 JLL. 3 4 0 3 . * -19?, 9. FRATIOICOFFF.1= _ _ f - P 303ICQE FF.1" S T O . F R R . c o r ' S T . " S T O . F R P . C O F F F . = S T D . ^ P R . pi 0.3357C-01 0.7430 0,) 779E 05 691.6 3753. kSQ = 0.3059 OUPPIN-WATSON STA." 1.136 _A!JTOC0°RELATION COE=F. " 0. 2694r.-01 _D-l_ 0.1443E 05* -303.3 DIP.. FRATIOI COEFF. )" 5 , 0 2 3 FPKnWICOEFF. )" 0 . Q T 3 9 STO. FPU. CONST. " ' , 9 5 ? , STO.FRR.COF.FF." 1 3 5 , 3 STD.FPR. 01 * I ' l l . •'SO = 0.2642 OUR RIN-W AT SO N STA." 1.472 AUTOCORPFI ATTON CQg=F. = 3.9040F.-01. Appendix 7. Coulter Counter data.Stenosomella ventricosa on laboratory food.E.S.O. method. One value omitted. 01 1717. 5421. 3114. 3896. 5378. 1612. 02 3.3 53 0E 06 -mmnA 0.41S4E 0.2 131E 06 J.2433E 06 0.3 57 5E 06 O.i Hit 06 C6 3 5 » 6 . 5904. _9_4 8 , 0_ 12 73. 527.0 _1_2.3_ 03 0.4246F 0.4802E 0.3C99E 0.3340E 0.3S42E 0.4P84F 04 06 3.000 8t- 4-.S88 06 3.000 06 2.000 06 3.000 06 4.000 06 5.000 2950. NAME MEANS 0.3 234E C6 0.3279E 0.5J76E 06 C.5C81E _0.1673E_06 0.1456F 6.3099S 06 0.2 768E d.le54E 06 0.1543E J L F . _ Q 6 _ 0 _ , 4 4 . 4 . 5 E_ 0 .2i '3E 06 0.2287E STO.DEV. D5 2.260 2.220 2.930 2.260 2. 320 3.120 2.340 l . ^ l O 06 3.280 3.280 3.280 1.410 1.410 1.410 1.140 1.140 07 0.9600 0.9600 0.9600 0.9400 0.9400 0.9400 0.9400 0.9400 08 8.000 A.000 8.000 8. 000 8.000 8.000 8.000 8.000 09 25.00 25.00 25.00 25.00 25.00 25.00 25.00 25.00 06 4.000 06 6.000 06 3.000 06 3.000 06 2.000 4& 7,£p_g_ n i 02 D4 05 _ P 6 _ 07 08 _P_-7675.31 311717. _ 2h 2 ? 12. 3.80 000 1 .90006 _2.09 20.3_ 0.900646 11.73)3 I 311.2 2 105957. II 46 5 7 . 06 5.000 CORRELATIONS m D Z _ 2.160 . 1.75 0 _0»°500_ 1.070 1.370 __.2tl.0_ 0.9400 Qj 1.140 1.130 .1,360. 1.940 5.850 _2*940_ 010 39.00 39.00 39.00 39.00 39.00 39.00 39.00 39.00 0.6700 0.9400 0.8200 _0.8 30O_ 6.7400 0.8900 _0,8800_ 010 3 7.2 000 1.42478 0.689944 3.9.090. 0.669184E-01 5.58654 1.44 7^0 1.0000 0.5626-0.5620-J i i - is. PS' 0.8300 OBSERVATIONS D6 0J_ 8. 000 20.00 22.00_ 22.00 13.00 J0,00_ 17. 66 25.00 26.00 25.00 25.00 30.00 _2B.00_ 26.00 39.00 36.00 36.00 36.00 22.00 .42,00 35.00 08 J 9 . 010 4. 57C0 8 0.2350 0.3392 -0,60 5__ 6. 08 2 4 -0.1164 -0.2689 1.0000 0."I 46' 0.7060' -0.0374 _0,36641 -6.0535 -0.0722 _ _ 0 8 _ _ 1.00QO 0.5355- 1.0000 0. 3138 -0.2532 •0. 3096 -0.291 8_ 1.0000 0.0294 1.0000 0.2869 -0.2083 0.7508' 0.1664 1.0000 •0.3836 0.1095 -0.7243--0.1503 -0.9434- 1.0000 •0.2479 0.1386 -0.3751 0.6772•-0.1966 0.1296 1.0000 0.2830 0 .O38 0.5799' 0.3687 0.3899 -0.5672. 0.3125 -0.3755 -0.7018* 1.0000 _ _ L • 0.9617E-C2* 02 5900. 4800, 5. 021 3.0278 FRATIOICOEFF.). Ft>903( COEFEjJi._ STO.FRR.CONST.' 12 96. STO.FRR. COEFF. * J.3U9E-02 _SJC i E_R. L . j ; =. 1554t 3700. RSO * 0.3165 OURBIN-WATSON STA.= 1.643 -AUT OC on R E.L AJ.I ON C3SFF. . 0.1409 1 1 260 0. • l S O O i 1 1 >. 1 _1 1 400.0 JLZ 0.1600E 06 O.30C0E 06 0.4400E 06 0.2300E 06 0.3700E 06 O.SIOOE 06 t o O Appendix 7. (Cont'd) r 5900. 1 I 1 > > 01 - -364.4 • 0.8878E-02* 03 •' 4800.. 1 FRATIOICOEFF.)- 4.002 FPROIUCOEFF.)- 0.0280 J • < STD.E R R.CCNST. - 1304. STO.ERR.COEFF.- 0.3t>24E-02 STO.FRR. 01 = 1555. 3700. 1 1 • • RSO » 0.3153 D'JKRIN-WATSON STJ.= 1.063 ..AUTOCORRELAT I ON C H F F . • -0.5735E-02 j i 1 , 1 • • • 2600. i • • 1500. ~~ at • • 1 1 1 -1 • 1 1 1 • 1 .1 1 400.0 1 . 1 0;llOOE 06 0.1900E 0.2 700F 06 06 0.3500E i 0.4300E 06 1 06 0.5100E 06 5900. 1 1 1 1 N 01 - 1540. • 298.8 • 04 4800. j . \ FRATIOICOEFF.)- 0.7597 ' | FP<OP.iqOEFF.|. J.4033 | STO.EF.R.CONST.' 1336. STO.EPR.COEFF.. 342.3 STO. P R R . 01 « 13?*. 3700. 1 1 1 < RSO =• 0.0552 0UR8IN-WATS0N STA." 1.657 ADTftCORRF LAT I ON C'H=«=. . 0.1653 1 1 • e • 2600. 1 >. • 1500. 1 1 1 I 1 1 1 1 '2 400.0 11 —I 2.000 3.000 4. 000 5.000 6.000 7.000 Appendix 7. (Cont'd) K : : 21 = 9BQ t? , » 891.9 » OS FRATIOICOEFF.>• . 1.691 FPR(J.R.ltQ.F.FF___5 0-2 14_ STO.ERR.CONST." 1381. STO.ERR.COEFF.- ' 685.9 STO.ERR. Dl 1768. RSO = 0.1151 DUR3IN-WATS0N STA.* 1.855 : AUTOCORRELATION C J E ' F . - 0.3649E-01 01 . 3118. • -37.75 « 08, FRATIOICOEFF.1- 0.1797 _FPRQj3JjaEFJ_.± -_ l ) . 6J ' 3_ STO.ERR.CONST.- 1153. STO.FPR.COEFF." 87.31 STO.ERR. 01 * 18 6 7. RSO =» 0.0136 DURBIN-WATSON STA. = 1.804 AUTOCORRELATION COEFF. - 0.81STE-01 JU • 4 3 3 6 . FRATIOICOEFF.I-_FPJ_18.( ,.COEF_L_i__ 7. 510 0«OL63_ STO.ERR.CONST.- 718.6 STO.ERR.COEFF.* 239.6 STD.ERR. 01 - 1496. _06_ RSQ = 0.3662 OURBIN-WATSON STA. * 2.428 AUTOCORRELATION COEFF. - -0 . 2266 FRATIOICOFFF.)* - F p ^ i l C J l f F F j . L i . STD.ERR.CPNST.-STO.ERR.COEFF.* 1. 013 -0-3343 4592. 334.3 H 1 Q . RSQ = 0.0723 OURBIN-WATSON STA. * 1.932 -LLT.QCORRELAT ION COf^E. - O. IS66F-0I _D__ 666.7 0.8886E--0_7..6.24_ FRATIOICOEFF.). _FP.R03XC0EF.F_J__ STO.ERR.CONST.- 6755. STO.ERR.COEFF.- 7481. STO.FRR. 01 . if 7 1 . 01 RSQ « 0.0068 DURBIN-WATSON STA.- 1, __UI.'iCaP-_£UI « OT 803 : 0. 83 50E-01 0-1 » -1497. » 117.2 FRATIOICOt e F.)a 1.13? FPROBICOFFF.I- 0.3077 STO.ERR.CONST.* 3749. STO.ERR.CrEFF.- 105.4 S T O . F R R . D l . 1303. 010 RSO * 0.0801 DURBIN-WATSON STA.* 1.958 AUTOCORRELATION COEFF. - 0. ie36E - 0 ? O Appendix 8. Coulter Counter data. Barnacle and Copepod nauplii on laboratory food. N.T.C. method. 01 02 03 04 05 06 07 D8 09 D10 V_ 433.0 J . 4 6 0 7 E 06 0 . 4 7 4 9 E 06 4. 000 1.200 2. 970 0.9500 13 .00 30.00 . 27.00 •< 104?. 0 . V V B E 0 6 o.? 5 1 0 E 06 2. 0 0 0" 1 . 2 9 0 1. 080 0 . 9 6 0 0 16 .00 28.00 42.00 9 ? R 5 . a. 4 4 1 4 = 06 0 . 3 6 0 7 = 06 6. 0 0 0 0.8-00 0.6400 0.9000 P. 000 25.00 74.00 M E A N S S T O . D E V . C O R R E L A T I O N S . 3 . O P S E P V A T I O N S 0 1 0 2 ' 0 3 04. 05 0 6 07 D8 0 9 0 ) 0 Dl 3 3 2 0 . 0 3 <• 3 0 4 . 4,3 1 . 0 0 0 0 ^ 6 6 * . 2 ! - - 0 . 7 3 0 1 1 . 0 0 0 0 J 01 4 S . 3 1 3 4 , 9 5 7 7 7 . 9 - 0 . 8 9 7 9 0 . 9 5 6 4 1 . 0 0 0 0 \ C4 4 . 0 0 0 0 3 2 . O O 3 0 O 0 . 8 4 1 3 - 0 . 9 8 3 6 - 0 . 9 9 3 4 1 . 0030 "5 .i_.ue.oo . 3 . 2 3 8 1 1 9 - 0 . 9 7 ! . 9 0 . 8 7 0 4 0 . 9 7 6 3 - 0 . 9 4 4 9 1 . 0 0 0 0 0 6 l . C 6 3 3 3 1 . 2 3 7 9 1 - 0 . 6 8 1 5 - 0 . 0 0 P 5 0 . 2 8 7 7 - 0 . 1 7 7 7 0 . 4 9 0 0 1 . 0 0 0 0 or 0 . 9 3 6 6 6 7 0 . 3 U 4 5 6 E -01 - p . 9 7 9 3 0 . 1 5 3 2 0 . 9 6 8 4 - 0 . 9 3 3 3 0 . 9 9 9 4 0 . 5 1 9 3 1.0000 09 1 ? . • 13 1 4 . 3 4 1 4 5 - 0 . 9 0 0 9 0 . ^ 4 7 " - 0 . op 17 0 . 9 8 2 0 0 . 3 1 6 5 0 .97*0 ! .0000 0^ r 7 , ~ S6"7 2 . 5 1 6 6 1. - 0 . ° 3 5 5 0 , - ' - ' ! . 6 0 . 6 3 ' 4 - 0 . 5 9 6 0 0 . 3 2 6 0 0. 3 9 6 1 0 . 8 4 ' 7 0 . 7 0 4 6 1 . 0 0 0 0 0 1 0 3 9 . 3 3 3 3 1 1 , 0 ! ' !. - 0 . 3 7 5 7 0 . T 0 7 4 0 . 7 4 5 4 - 0 . 8 1 7 1 0 . 5 8 3 3 - 0 . 4 2 2 2 0.5554 0 . 7 2 6 3 0 . 0 2 4 0 1. 0000 r \ ni » 0.5450P 05«~0 .109T * 02 3300. 6700. X \ J ( FRf.TI0tCnF.rP.)* 1.141 F P ' . P S M C O E F F . U o.v r ^5 1 >. I « 1 » •s S T I > . E R R . C O N S T . ' 0.47965 05 STO.K*R.Cf< rFF.' 0.1327 -S T P . = " > . M = • f i j ) 3 . 5100. 1 • " S O • 0.5330 0U39IN-WKT*ON S T A . ' 1.325 A'JT Of OF RF L ^  T I ON C T : = F . ' -0.7514E-01 I • 1 • 1 • 3500. 1900. 1 • 1 • I . I . I . 1 300.0 1 1 1 1 1 1- 1 -—1 0.4380E 06 0.4620E 06 0.4860E 06 0.4500E 06 0.4740E 06 0.4°BOE 06 I Appendix 8. (Cont'd) r 8300. - 1 1 > 1 • \ ni = a.7}oy. o 5 » - o , 4 0 A 5 F - a i * rn 6700. 1 • FRAT10(CHEFF.). 4.153 ....F = M»(CO = FF.)= 0,7 -vol 1 0 1 • t • < STI1.ES».C1SST.» 3276. S T 0 . c o 9 . C N E F F . B 0.1.F79E-01 5Tr,.=r3. HI T. ? S 3 3 . 5J00. - • • It *?0 = 0.3361 ni|l'3!N-WAT70N S f A, = 1.711 A U T C C O S O F I i ' M N c.o===. « -0 , ?»TO- j • 9 i 3500, j '•' 1900. 1 • • • — • 1 . 1 3 00.0 1 e 0.3560E 06 0.3950E 0.4340F 06 \ 06 0.4730F 0.5120E 06 06 0.551 OE 06 • 8300. j 1 \ P I * -392?. * m i l . * 04 - 6700,. F ^ ^ T I C I C O F F F . ) . 2,422 ..e?.330<.C_FcF. j , 0.3677 j i ST0,5PP.C0*iST.- 5327. STO.EPP.Ci'FF., 1164. S T ^ . r - B . 01 T 1-XJI. 5100. 1 PSO = 0.7.378 0'.T3P:---IATSP'I STA." 1.500 .... M'TOO'RCLATT-w C ' 3 . E B . » -0.1667 ! » • * 3500. j • 1900. I • f « ——— 11 -3 00. 0 ( « 1 2. 000 2.800 3. 600 4.400 5. 200 6 .000 Appendix 8. (Cont'd) J J J « 0 . 7 7 8 7 * Q 5 - 0 . 1 7 5 7 F C 5 * 05 F R A T I O ( C O E F F . ) • -FnMQ.(_nEFF,Jj S T M . f R R . C O N S T . * S T O . F R R . C O E F c . ' S T O . ^ R P , I l l 1 7 . 0 4 _0.1.T3__ 47 3 6 . 4? 5 4. 14 3* P.SO = 0 . 0 4 4 6 O I P 3 I N - W A T S 0 N S T A , ' 7 . 1 4 ? .. A I J T o r p . r . p c L A T 1 0 1 C T - ' F . - - 0 . 3 8 10 FRATI0(C0EFF.1= JFJ>PQ1_C_.-F. F.l» 4 . 8 1 1 0 .781 .4 S T D . F R R . C O N S T . = 5 4 4 1 . S T O . F R R . C O r F F . 3 4 4 1 . 9 S T O . m a . 01 » P S ' J - 0 . 1 7 79 D U R B I N - W A T S O N S T A . = 1 . 7 6 5 - . A U T O C O R R E L A T I O N C O E - F . . - 0 . 2551 •2370. _ D _ _ F R A T I O I C O E F F . 1- 0 . 3 5 7 1 ... F . " * Q . ! U C O E F F _ _ _ 3.5,12j_ S T O . E R R . C O N S T . = 4 7 3 8 . S T O . E R R . C O F F F . - 2 f 4 5 . S T O . F R Q . 0 ) ' 4 4 5 5 , PSO = 0 , 4 6 44 OIMHIN-WATSON. S T A , ' 2 . 7 7 1 . AUT3C - 1 R R E LAT ION C ' l . - ' P . = - 0 . 59 0 4 J _ _ 0 . 4 7 5 9 F 0 5 * - 1 6 0 0 . « 00 F R A T I O ( C O E F F . ) = 7 .011 turn HJLCQ££ r . i . 0.7475 S T D . F R R . C O N S T . = 0 . 1 . 6 7 7 E S T O . F R R . COFF>=. ' 4 0 4 , 3 S T O - F R P . 01 ' 7| « . i . 05 PSO = O.E 752 OUR 31 N-W A T S f N S T A , ' 2 . 9 7 4 •AUXQCQgjR F.LAT . I ON C O E ' F . ' - 0 „ 6 1 13_ n i ' 0.1 7 4 2 F 0 6 * - 3 . 1 3 1 I F C4 » 07 01 i 76 7 7. + - 1 4^,. R # n i n F P A T I O I C O E F F . )- .21.44 . (.CO EF F ..1 ~ _ 0 . 1 5 5 S F R A T ! Q ( C 0 E F F . | * F P a o n i r . Q F F r . ) ' 0 . U 4 4 0 . 7 4 1 4 S T O . E R R . C O N S T . ' 0 . 2 5 3 3 E 05 S T U . F R K . C O C F F . . 0 .2 7 0 9 6 05 S T O . F R O . oi = l 7 i 2 . S T O . F R O . C O N S T . ' S T O , E » P . C O E F F . ' C T n . r o o . n i 0 . U 1 1 E 05 3 5 7 . 2 PSO » 0 . 9 5 9 1 OUR 8 IN - w A TSP N S T A . ' 2 . 2 1 0 A U T O C C R R E L A T I O N C n ' - : - e , ' - 0 . 4 0 3 ' PSO = 0 . 1 4 1 2 DI IR9IN-WATSC-N S T A , = 1 . 0 1 6 S l l T n r n o o F i AT 1HM r n c c c - n c / . o c c n i Appendix 9. Coulter. Counter data. Barnacle and Copepod nauplii on laboratory food. E.S.O. method. ( 01 02 03 04 05 06 . 07 08 09 010 s ? V ! 4'. 3c 06 0.1432E 06 4, 000 1.110 7 .970 0.9 500 13 .00 30.00 27.00 i 1515. 3.1 S * I F Ob 0.1 7 ( ,6F 06 2. 0 0 0 1. 080 "1.080 0.9600 16 . C O 28.00 47.00 23 r 8 . 3.! 39 5 0 0 4 0.13 70= 06 2.000 0.) 903 0. 1900 0.P30O 17 . 0 0 26.00 35,00 3) :-3. 3. 3 32 7 F 05 0.2H71E 0 5 3. 00 0 0.4100 0. 4100 0.9100 8. 000 25.00 74,00 9 ; ' 0 i . 3.4207F 0 6 0.3457E 06 6. 000 0.6400 0. 6400 0.9000 9 . 000 25.00 74.00 Nt«F MEANS STO.OFV. CORRELATIONS 5. OBSERVATIONS \ 0 ] "2 03 04 D5 06 07 03 09 010 Dl 3 4 0 7 , - 3 3653 .77 1.0000 < 02 . 1>"!3 70, 1. 4 J. 4 26. 0 . 8 / 9 ! . 1.0000 03 l c ^ o , 1 1 4 4 4 % .0. 7 45 5.. _.0.9F .7 8. J,0000 04 3. 4? v l O l 1 . 4 ? ' 3- o ; 7 8 ' s 0.7943' 0,7 o n 1.0000 15 0 . 6 = 6003 0 , 4 )ST ) 2 -0 .7766 0.;53? 0. 21 P 7 0.147 8 ).0000 14 l.J<?03 1 . 1 ! - ? l -3„ <>2 6 6 -0.006? 0, 0 1 0 1 0 , 2 0 0 9 0 . 7 9 7 ) 1.0000 07 n. i i 2oi) 0 . f»J 4 7 = 1 F -01 -0.2512 -0,3203 0. 0!<- 5 0.!141 0.9187. 0.6565 1.0000 0 9 1.2,4 3 0 3 4.2? 7B 5 - 0 . 6 4 ) 6 -o.r84?. - 0. 1 49? -0.7264 0.1143 0.)005 -0.1362 1.0000 _ 2 * . 9>V)0 2 , ? 9 5 - 0 . 6 7 5 0 -0.175?. -0. 058 J - 0 . 1 . 7 9 2 Oo 7Q70 0.9029 0,6049 0.41^0 1 .0000 O 1 0 2=.-333 9.70631 -0 . 3 2 38 - d . ' Y i ' j -0,0?] 4 -0.7173 0.0557 -0.3331 -0 . 0235 0.7666 0. 0b<.« 1.0000 9 8 0 0 . 1 1 n i x - 4 2 2 . 3 • 1 . . ? l ' . t = - o j + 0 2 7 9 0 0 , j F R A T I - H C O E T . l . 4 , 1 1 6 F P ^ n ( C i E E F . I = 0 . 3 - > 3 9 j < S T O . U P o . r . M N S T . . 1 ) 1 9 . S T O . ?=.? . . C O E F F , = 3 , f 5 5 0 E - 0 2 S T i l . r u , n i « .", 1 ' 1 , 6 0 0 0 , • RS 3 = 3 . 5 7 0 9 r>'Jt>-3:>!-w.MSr'l S T A . » 3 , 7 4 1 4 f . l * O C . i ' ' . - T l * T I J N : - > " F . » 3 , 4 1 . 1 4 j 4 1 0 0 . [ 7 7 0 0 . 1 1 * 1 . ° 1 3 0 0 . 0 I # l - l — 1 — i 1 1- 1 1 0 . 2 0 0 0 E 0 5 0 . 1 0 0 0 E 0 . 1 8 0 0 F 0 6 0 . 3 4 0 0 E 0 6 0 6 O . 2 4 0 0 E 0 6 0 . 4 7 0 0 E 0 6 ' Appendix 9. (Cont'd) 9 8 0 0 . j \ 01 = - 4 7 9 . ? * P . 2 3 7 8 E - 0 1 * 0 3 7 9 0 0 . ' < FRATmCOEFF. !» 3 , 752 F P R 0 8 1 C ^ E F F . ) = 3 , ! 4 » i • • < S T O . f » R . C O t v S T . = .3407. S T 0 , F ^ P . C O F < : ' : . = 0 . 1 2 3 9 E - 0 1 S T O . r o n . 01 - 71 T O . t ooo . • PSO = 3 , 5 5 5 7 OURRT'I -WATSON ST 1 * . « 0 . 6 3 6 3 4'.1T 0<" n c R •= |_ s T ln\J C 3 : C F . « 3 , ^ 4 8 8 j • • 4 1 0 0 . j C « 2 2 0 0 . | X j « • 1 3 0 0 . 0 • 1 - 0 . 0 0 . 7 0 0 0 E 0 . 1 4 0 0 E 05 06 0-2800E 0.2100F 06 06 0 . 3 5 0 O E 04 f j 9 8 0 0 . I 1 U- 0 ' = - 2 1 1 ' . , • 171.0. * 04 7 9 0 0 . ! 0 j F R . I T I O I C O E F F , ) * 4.77,3 c p j n t i i f n t t c , |. o . l \ 7? j • < STO.FRR.CONST. • 3? 16. <Tt>.r.SK.CO?rF.» 7 1 . 3 . 3 S T O . e - J o . n i . 3 6 1 8 . 6 0 0 0 . -• C 5 ) = 0,>,143 r>l1"8:y-w.\TSt'N STH,= 1 . 1 0 0 MIT r . l R R F l A T l C N : 0 : : c . » - 0 . 3 1 5 9 5 . - 0 ? • * 4 1 0 0 . j • • 2 20 0 . I 1 1 . • 0 1 1 . 1 > m 3 0 0 . 0 I— . . . 1 | — i 1 — | — 1 2. 0 0 0 2 . 8 0 0 3 . 6 0 0 5. 2 0 0 4 . 4 0 0 6 . 0 0 0 J Appendix 9. (Cont'd) ni . •5704. • -74 87. * 05 ni • o.io?9= 0 5 * -547.5 * 0 3 > FOATI0(C0F c F.1' FPRO 3 ( 0 0 F F F . l > 0.2486 0.6517 FRAT^0(COEFF.1' FP»n9( C O E F F . | = 2, 0 9 9 0.7 4 34 S T D . E = . R . C 0 M S T . = S T O , F P R . C O E F F . ' S T O . F O ? . 01 3P 72. 4? 80. M M . S T O . F R O . CONST.' S T O . E R R . C O E F F . ' STO.FRR. 01 ',7 04. 377.0 3-> 34. PSO - 0.0765 01IR8IN-WATSON S T A . ' 1 . 1 5 0 AUT0C1RRFI AT ION C ' ) .= F . * 0 . 1 3 8 2 F - 0 2 RSO = 0 . 4 1 1 6 DUR3IN-WATSCN S T A . ' 2 .250 AUTOCOR P r 1. AT ION C O " F „ = -0 .4702 n) = 4971. • - 1 3 3 ? . * 06 01 = 0.3396F. 05* -1.1 3 7 . * 09 FPATin(COEFF.1-F n - J O ^ C O F ^ F . ) ' 0.6 675 0, FRATIOICOEFF.)' FPR03(COEFF.)= 2,511 0 . 2 1 1 2 STI.l.PRR.CONST. ' S T 0 , E R R . C f 1 F F F . = S T O . ' o o . ni 248 2. 1704. 38 1 3, STO.ERR.COMST.= STO.EPR.COEFF.' S T O . C C R . 01 0.19.27E 05 71 7.3 31 1 0 . F.SO = OUR3fN-WATSON S AllT'lCORPFLAT I ON 0.13 20 TA. » 1.220 CO£=F. . 0, 12 9 3 E - 0 1 RSO ' 0.4557 DUP P.IN-WATSON STA,= 1.675 AIITOrOPOFl .AT ION C02FF, =, -0.1611 01 . 0 . 1 P 7 1 F 05<-0.1 737-= 054 07 01 • 7575. • - 1 3 3 . 7 * 0 1 0 1 FPATIOICOEFF.)' F P R N 0 ( COF P - F « > » 0 . 2 0 2 ) 0 . 5 8 1 ' F°ATIO(COEEF.)a FPP.ORI COEFF. 1» 0.3685 0.3PR3 STO.E^S.CONST. = S T O . F P R . C O C F F . = S T O . c o = . 01 0. 1611 E 05 0.3943E 05 40 RO. STO, E " P . CONST. -STO.FRIJ .COPFF. ' S T O . F R R . 01. 4940. 22 3 . 5 37 ' 8 . R S O « 0 .06?! 0UP.9IN-WATS0N STA.= 1.002 AMTOrnoo.F I AT TON C : V " F . ' 0.7539F-01 SSO ' 0.10=54 0UR9IN-WATSnN STA. = 1.234 A1IT0C0PPFLAT ION CO?=F. = -0.1020 | Appendix 10. Coulter Counter data. A l l samples. N.T.C. method. oi 3334. 0.1640F 05 3449. 0. K>70E 05 43A .O 1351. 11?4.. 0? J.33J3E 0.8316E 0.3 3> 4E O.4340E 0.4330F i>. T.? •» 0 F Appendix 10. (Cont'd) 5972. 633.0 1042. B?85. 915.0 NAME MEANS 0.64416 06 0.6012E 0.4607E 06 0.4749F 0.4T79E 06 0.5510= 0.4414F 06 0.3607E 0.5121E 06 0.3766F STn.OEV. 06 3.000 06 4.000 06 2.000 06 6.000 06 4.000 CORRELATIONS 0.9500 0.2200 ' 0.5600 18.00 26.00 1.200 2.970 0.9500 13.00 30.00 1.290 1.080 0.9600 - 16.00 28.00 0.B400 0.6400 0.9000 8.000 25.30 0.5900 1.740 0.8500 10.00 25.00 66. OBSERVATIONS  20.00 22.00 47.00 24.00 39.00 01 __2_ 5633.72 57972*. 8390.22 349514., 03 04 536900, 5.43 9 H .1 i?7 2_L6_ Dl 1.0000 0.1684 02 1.0000 D3 04 05 06 07 09 mo 3335^9. 2.12)43 _0__4J>?0hS 04 1.61377 07 0. r l j o i ' j ..08 9,40003 09 22.6404 010 42.6816 0.0603 0.4618 0.1175 1.56421 0. < 9194 RE -01 _3.01778 5. )•> >1 7 17.816? 0.9494 •0.2415 •0.7871 -0.3633 0.0370 -0.1439 -6.299 8" 0.001 0 1.0000 -0.3719 1.0000 TO. 1474 0.2559 -0.0525 0.1094 0. ?.R54_ : 0 . 31 1 6 •0.7719 1.0000 -0.0656 -O;093O 0.1 752 -0.0733 0. 295 3_-;0. 3459_ -6. 123 4 -0.1 756 -0. 252 5 0.7738 0.2466 1.0000 0.2454 ; 0.2224 1.0000 •0. 2977_-O.O28 0 -0.) 87 3 1.0000 0.1534 -6.1517 -0.068 6 0.704T" 0.1808 -0.2041 0.2492 -0.76)1 'lTOOOO 0.0739 1.0000 _DJ_. 328 1. » 0.40558-02* 02 0.44OOF 05-0.3500E 05 FRATIOICOEFF.). 1.869 — F W U . C O £ F F . | « 0.1 7?9 STD.ERR.CONST.. 2002. S T O . f R O . C O H F F . . 0.2966E-02 S T O . C R O . Q l , ^ I S . 0.26 OOF 05 RSO » 0.3204 0IIR8IN-WATS0N STA.= 0.8165 ..A'JT JC'JiSELATIo__C0^ ;F. . g. «,ft«7 0.1700E 05 8000. 1 11 21 • 3 21 1 2 211 11 . 1 221 -1000. 211) 252 22)2 0.9000E.05 0.8300E~06 0.1570E 0? 0.4600E 06 0.1200E 07 0.1940E OT Appendix 10. (Cont'd) T C I ? . • 0 .1SlTF-rw« FRATIOICOEFF.). (1.3336 _E£ROa tiQ EFJ=__s__3_6J_7_ STO.ERR.CONST.* 1179. STO.FRR.COFFF.. 0.3118E-0' STO.FRR. ni » l ^ H , RSO = 0.1036 0UR8IN-WATSON STA.* 0.9119 _AUT0C2R.RELAIl:JN_C0; r i :. = 3.8912 0.3S00E OS 0.26 OOF 05 0.17 OOF 05 -1000. - 1 L . 1 ? 12 1 1 .1 > 3221 1 12213 21 11 1 14 21 212 2 3) 21 0.1200E 06 0.8400F 06 '0.15606 07" 0.4800= 06 0.1200E 07 0.1970E 07 _Q1 » -4306. -1827. « 0 4 FRATIOICOEFF. )• 17.35 FPRQBtCOEFF. j . Q.aoO]_ STO.FFP.CONST,« STO.EPR.COEFF 2559. 43 3.7 . 7100. RS3 » 0.>133 DI)P8IN-HATSrN STA.» 0.7712 -AUTOCORRELATION C J t=F, . p . 4 I ^ T 0.35 OOF 05 0.26 00E 05 0. 1700F 05 219 f t er i • H U1 i a <1. o q u u U <J • • — —1 OC CC ( c ccl or or t . <- O U J i n 1 I- cd • • < a j o o ! ac UJ(- 1- > U. VI trt t or t - i . r - i U- U. f U. I u i d c c q o q u o <_> >J • • — ac ar orl oc a o J c c ~ o ar <t a ct u. c a d • • tu > • c < o * -v. 3T C r 2r — I or 0> .-i < •a tr- \r c o c a P. <3 i n tn c u Cr U" c 9* r O l> i n u | 1- c t 1  HI R n nl c q u o g or < OJ O of i j a cr cc cr UJ U-• « o c t -trt * H I M > o t-j : 00 < : t - -i Sal in r .<t|. u. u J ; J tr, u . c c 3 O O • • w oc a cd C g ct or ed < a l o o d • B tU "> • c e? -z -i/> c » iXL t / i -a. u. 1 t « i/i • tn tr» c | -; O J o tr c 1 0 <-S| 1 • n . . V- u_ C C C I oc a crl c 7 ~\ 01 u". < 11 I Appendix 11. Coulter Counter data. A l l samples. E.S.O. method. 0? 03 0 4 D= 06 0 7 08 07 ; ni 0 v ??•»* . 0 , 4 7 5 7 = . 0 6 0 . 3 3 0 6 = 06 4 .000 0 . 5 9 7 0 0 . 4 8 0 0 0 . 7 7 0 0 ".. 000 1 2 . 5 0 « 6 . 00 J f 0 . 1 6 4 3 8 05 0 . ,9 >?. 3 = 0 6 0.7 3 7 6 F 0 5 8 . 0 0 0 0. 8 5 0 0 0.4 8 0 0 0 . 7 7 0 0 9, 000 1 7 . 5 0 4 6 , 0 0 < • H •'•••>. 0 , 7 1 1 7 = 0 6 0 . 5 0 7 1 = 0 6 6 . 0 0 0 0 , 6 5 0 0 0 . 7 8 0 0 0 . 7 7 0 0 9, 000 1 7 . 5 0 46.00 1 0 - 1 ? 7 0 = 0 5 0 , 3 3 \ 4F 0 6 0 . 5 3 6 5 = 3 6 9. 00 0 0 . 8 5 0 0 0 . 7 8 0 0 0 . 7 7 0 0 " 9, 000 1 2 , 5 0 4 6 , 0 0 ! 1 7 1 5 , 0 , 1 . j V 7 = 0 6 0 . 1 4 7 7= 0 6 2, 0 0 0 1 . 3 5 0 1 . 8 9 0 0 . 7 7 0 0 1 0, 0 0 1 5.30 ?4., 3 0 I 7 5 74. 0 . - 7 H 4 = Oft 0 . 2 0 4 7= 0 6 4 , 00 0 1 , 4 = 0 1 , 7 9 0 0 . 8 1 0 0 1 0 - 0 0 1 5 , 0 0 3 4 , 0 0 V ' " 0 . 0 0 , - 7 7 - . = 0 5 0 , 7 1 5 1 r 0 5 1 . 0 0 0 ' 1 , 2 1 0 0 . . 9 2 0 0 0 , 8 9 0 0 1 0 , 0 0 1 5 . 0 0 ?4 00 • 1 " : . 0 3 13 5 = 0 6 0 , 2 4 3 6 " 0 6 3,. 0 0 0 1.01 0 1. - 4 7 0 0 , 9 6 0 0 1 0 , 00 1 5 , 3 0 34 no < l o 0 , 9 ) ? 1 C 0 5 0.6 3 5 0 = 0 5 2 . 0 0 0 0 . 8 3 0 0 5 , 3 0 0 0 .8 7 0 0 CT,000 1 6 , 3 0 7 9 , 00 . 1 . 6 ."A.7=. _ . - < 7 _ O, 5 4 j _ , r 0 6 9„ T O O 1 . ' - 9 0 0 . 7 9 0 0 0 . 9 7 0 0 P, 0 0 0 1 5 . 0 0 4 7. 00 0 , 4 4 1 9 = 0 6 0 , 5 9 , 7 ? 3 6 0 , 5 4 1 4F 0 6 9„0OO 1 . 7 2 0 0 . 2 8 0 0 . 0 , 8 7 0 0 8 , 0 0 0 1°, 3 0 4 2 , 30 0 5 9 , ,V)'. r> = 0 6 0 , 4 9 6 6= 0 6 8 , 0 0 0 1,4 = 0 0.5 3 00 1 , 0 0 0 8 , 0 0 0 ! >7 . 0 0 4 7.00 0 , ?.7* 1 = 0 5_ _o , ( 5 ' . <j = . .06 _ _ 0 , 4 = f T . Q 6 _ _ 8 . 0 0 0 1 . 7 7 0 0 , 57.00 1 . 0 0 0 8 . 0 0 0 1 9 - 0 0 47. 00 1 V . 7. :'). >'. > >. F 0 6 ~b. 4 i s 5= 0 6 8 , 0 0 0 1 . 4 5 0 4 . 1 1 0 0 , 9 7 0 0 8 , 0 0 0 1 3 . J O 4 2 . 0 0 0 • 6 1 1 1 ~ 0 6 0 . 4 i n 5= 0 6 8., 0 0 0 1 ,77 0 4 , 1 1 0 ; 0 . 8 7 0 O P., 0 0 0 1 9 , 0 0 4 7 , 0 0 1' 4 . 1 ! «F_ . 0 6 - • 0 . 3 6 4 7 = 0 ^ 7. 0 0 0 1 , 4 = 0 5. 6 8 0 0 , 8 7 0 0 p , 0 0 0 1 3 . 3 0 4 ? „ 0 0 ! 1. >v. > 1 i = 0 6 0 ~ 3 (• •'•7= Oft 7., 0 0 0 1 . 7 - 0 5 . 6 5 0 0.. " 7 0 0 H , 0 0 0 1 M. DO 7, 0 0 1 . 5 0 6 . 0 , 4 7 3 ? = C 6 0 . 3 F 6 2 F 0 6 7 . 0 0 0 1 . 4 9 0 4. 8 4 0 0 « 8 6 0 0 8 , 0 0 0 1 8 . 0 0 47-. 00 0 6 0 , 1-1.42F 0 6 7., 0 0 0 1 , 77 0 4 . 3 4 0 O . " 9 0 0 P, 0 0 0 1 p . 0 0 4 ? 3 0 74 7 4 . 0 , 4 7 7 7 = 0 6 0 . ? 5 0 6 F 0 6 9 , 0 0 0 1 . 4 0 0 2 . 7 1 . 0 0 , 91 0 0 A, 0 0 0 1 P. 0 0 . 4 7 , 0 0 0-4 2 7 7^ 0 6 0 . ? >36'7 0 6 7. 0 0 0 1. 7 7 0 2 . 7 1 0 0 , 9 1 0 0 9 . 0 0 0 1 P . 0 0 4 2 , 0 0 ____ o._ ,V. M H F 0 5 0 6 1 , 0 0 0 ' 1 , 0? 0 0 - 7 7 0 0 0 , 8 7 0 0 P. , 0 0 0 1 5 , 0 0 40, 0 0 "6\'->v '?"" 0 6 0 , 1 977=" 0 6 5 , 0 0 0 0 . 9 1 0 0 0 , 2 0 0 0 0 , 9 7 0 0 ' >'s"dob"' I P , 0 0 4 9, 0 0 ) " • • > . OA 0.1 M 5 r 0 6 5 . 0 0 0 0, 1 6 0 0 0 . 7 1 00 0..OI 00 5. 0 0 0 1 9 . 00 '••>. 30 t.™7. 0 0 , ' 1=7 = 0 6 0 , 1 0 7 7 = 0 6 2 - 0 0 0 1 . 1 3 0 0 . 6 4 0 0 O . B 6 0 0 9 , 0 0 0 1. P . 0 0 9 0 : 0 0 51 - 8 . .)., " > •- •T- 0 . 2 7 6 4 = 0 6 H„ (IC 0 2. 0<- 0 0 . C 4 O 0 0 . 8 6 0 0 8„ ono 77 .00 " 0 0 0 3 9 3 a , 0 , ,7 2 1 3 r 36 0«.??' .4F 0 6 8 * 0 0 0 1 . 3 8 0 0, 6 4 0 0 0 , 8 6 0 0 8 , 0 0 0 2 7 . 0 0 9 0 , 0 0 .<•:•:. !>. 0 . 1 " OS 0.1 v ) 7 i : 0 6 4. 0 0 0 1 . 0 0 0 0 . 4 2 0 0 0. 6 6 0 0 H , 0 0 0 2 7 . 0 0 7.6,00 9 5 5 5 , 3 = 7 1 7 1 = 0 4 0 . 2 9 1 . 6 = 0 6 e. 000 1 . 4 7 0 0 . 4 3 0 0 0 . 6 6 0 0 8 , 0 0 0 2 7 , 0 0 ?6 ; 3 0 5 3 4 3 . 1 )7S = 0 6 o.roi5= 0 6 8 . 0 0 0 1 . 1 7 0 0 . 4 2 0 0 0 . 6 ' . 0 0 8 , 0 0 0 2 7 . 0 0 3 6, 0 0 " 0 6 . i . .7 i r ^ 1 - 0 6 0 , 3 5 0 T 0 6 6„ 0 0 0 1 . 0 6 0 0 . 7 9 0 0 0 . 6 6 0 0 1 « , 0 0 0 7 7 , 0 0 3 6 0 0 0,1 76 1 F ~ 0 5 ~ "oYi"i"?' vr 0 6 0 . 3 5 9 7F. 0 6 8 , 0 0 0 1 . 4 7 0 0 . 3 9 0 0 0 . 6 6 0 0 8 , 0 0 0 7 U 3 0 3 6 , 0 0 p 7 9 7 . o, i 17 irj 0 6 0 . 7 . 5 9 7= 0 6 • 7.. 0 0 0 1 , 1 7 0 0„ 3 9 0 0 0 ,61 0 0 5, 0 0 0 77, 0 0 2 6 . 0 0 1 7 0 7 . 0 ,!. ' i •> = 0 6 0 , 1 7 1 7 = 0 6 3, 0 0 0 1 . 1 5 0 0 , 4 8 0 9 0 . 8 7 0 0 8, C O O 2 7 . 0 0 =-0,00 6 4 1 6 . 1 . 2 7 •. .1 -. 0 6 0,r .'a 6 = O o 6, 00 0 1 . 57 0 0 , 4 P . 0 0 0 . " 7 0 0 9 , 0 0 0 7 7 , 0 0 9 0 - 0 0 7 9 < M . 0 , 2 1 F 0 6 0 . 7 9 7 4 = 0 6 7. 0 0 0 7 = 0 4 0 Oc 4 8 0 0 ( 3 . 8 7 0 0 8 . 0 0 0 2 7 . 0 0 0 0 . 0 0 0 . 2 4 3 6 = 0 6 o.r .771 = 0 6 6, 0 0 0 1 . 0 0 0 0 . 7 6 0 0 0 , 9 ? 0 0 9 , 0 n 0 2 7 , 3 0 3 4 . 0 0 6 4 9 7 . 0 . 3 1 ! 4 5 0ft 0 . 7 9 7 0 = 0 6 8 , 0 0 0 1 . 4 7 0 0 . 7 6 0 0 0 , 3 1 0 0 8 . 0 0 0 2 7 . 0 0 "6 , 0 0 0. , H I 4 = 0 6 3.?17 0= 0 6 P . 0 0 0 1 . 1 7 0 0 . 7 6 0 0 0 . 8 1 0 0 8 , 0 0 0 77.00 ?ft,00 4 6 1 . 9 0 , 5 3 1 9 = 0 5 .0.5 2 5 3 = 0 5 2 , 0 0 0 1 , 1 7 0 1 . O ' O 0 , 1 7 0 0 R, 0 0 0 77 .00 n o , 0 0 0 . 1 7 V I F 0«- O . i 91 5F 0 6 6, 0 0 0 I . O O " 1 . 0 3 0 0 . 9 3 0 0 P. 0 0 0 ? / „ 0 0 9 0 . 0 0 ; -.'7 7 . 0 . 71.5 3 = 0 6 0 . 7 7 2- ? F 0 6 7. ono 2 . 0 4 0 i . O l O 0 . 9 2 0 0 8 , OOO ? 7 „ 0 0 ° 0 : 3 0 . ) . ' 7 4 1 = 0 6 0 . ' 1 76 = 0 6 4 . 0 J 0 1 . 7 7 0 0 . 6 1 0 0 . O . 6 1 0 0 1 5 - 0 0 7 8 , 0 0 7 0 , 0 0 l a w . . ^ •jl> 0.1 : o 7 r 0 6 M M 0. ; ) 7 O 0 0 . 6 7 00 0 . 6 9 0 0 : 5 , 0 0 2I>. OO ?'K 1)0 0 . 7 ' . 7 0 f 0 5 0 , 1 I 7 > = 0? 0 . 1 01 OF 0 7 4,. 0 0 0 0, 9 4 0 0 0 , 4 7 0 0 0 . 9 6 0 0 9 . 0 0 0 1 5 ., 5 0 5 7 , 0 0 0 , 1 t i n e 0 5 r\ . 0 7 ) 5 F 0 6 0.6 J7/»r- 0 6 6„ 0 0 0 1 . 0 0 0 0 . 6 4 0 0 0 . 5 8 0 0 1 0 . 0 0 ) r , 0 0 ' 4 , 0 0 <••!>?. 0 . 2 t t ' t l" ~0ft 0 . 1 4 ) •)=" 0 6 4, 0 0 0 0, 61 0 0 0 . 7 9 0 0 O . r . ' f . ' i ').. 0 0 0 ? ' . 0 0 . v . . 00 1 = 0 0 . 0 . 1 1 4 1 " 0 6 0 . 4 7 5 1 = 0 6 8 . 0 0 0 7. P R O 1 . 3 90 0. ° 0 0 0 1 0 , 0 0 7 6 , 5 0 7 0 , 0 0 °7 7. 0 0. 7-3? .5 = 0 6 0 . 1 7 9 3= 0 6 2 . 0 0 0 0. 5 4 0 0 1 . ? " 0 0 . 9 6 0 0 1 0 . 0 0 1 5 . 5 0 3 0 , 00 1 1 - 0 . 0,1 7 4 6C 0 6 0, ' . 16 4= 0 6 6 , 0 0 0 7 , 9 3 0 2 . 5 8 0 C . 6 9 0 0 9 , oort 7 4 , OO 1 6 7 . 0 3. 1 0 9 917 0 5 0 . 1 217C 0 5 !., 0 0 0 3 , 1 1 0 1 . 6 0 0 0 . 6 9 0 0 p , 0 0 0 2 4 . 0 0 ' 6 . 3 0 ' 0 6 . O., 7 5 7 an 0 5 0 . 7 0 3 4 c 0 5 ? 0 0 0 0 j . 6 9 0 0 . Q ? 0 0 0 , 7 2 0 0 p . 0 0 0 7 ' . 0 0 3 6 , 00 i •»: s . 0 . 3 .V) 1 = "Of. 0 . ? ) - ' t O c 0 6 5. 0 0 0 1 . I^O 1 , 4 5 0 0 , 7 2 0 0 ) 5 . 0 0 ? ' , 3 0 7 7 , . 1 0 7 i 7 . 0 0., V 17 0 " OA 0 „ ? 573 = 0 6 3 = 0 0 0 2 . ? ? 0 5 . 3 9 0 0 , 6 8 0 0 1 5 . 0 0 2 9 . 0 0 77 , 0 0 ! ".,). 0 •3.1 42 4" 0 6 0 . 1 57 l.r 0 6 7, 0 0 0 1 , 4 5 0 4 , 5 5 0 0 . 6 F . 0 0 9 , 0 0 0 15. 5 0 = 7. 0 0 4 7 0 . 0 0.1 !>'. = 0 6 O . i . 7 ' 4 F 0 6 2 . 0 0 0 0. 8 7 0 0 0 . 5 5 0 0 0 . 7 7 0 0 9 , 0 0 0 i * - . 5 0 57. ?0 " " 6 7 . 0 .1 , ? 31 5 = 0 5 0 . 7 666 = 0 5 1..00C 1 . 6 2 0 0 . 5 2 0 0 0 . 7 8 0 0 9 . 0 0 0 1 5 , 5 0 8 7 . 00 5»>'7. 0 0 , 4 .'6 7 = .15 0 , 3 7 74= 0 5 3 . 0 0 0 1,7-0 • 2 . 7 3 0 0 , 8 0 0 0 8 . 0 0 0 7 5 . 0 0 . 7 4 , 0 0 i q 2 7 « •7 , 4 3 7 3 = 0 6 0 . 3 7 4 7 = 0 6 6; 0 0 0 0„ 9 4 0 0 1 c 8 5 0 0 . 7 5 0 0 p , 0 0 0 7 5 . , 0 0 2 4 - 0 0 ! 7! 7. 3.. 3 33 0 = 0 6 0 , 4 7 4 6 = 0 6 3 . 0 0 0 7 . 2 6 0 3. 2 8 0 0 . 1 6 0 0 Po 0 0 0 7 5.,00 3 9 , 3 0 V. " 1 6 7 , 0 ) , " 1 3 9 = 0 6 0 , ? ' 7 5 9 F 0 6 3-or.O ? . ? 7 0 3. 2 8 0 0 . 9 6 0 0 8 , 0 0 0 • 2 5 . 0 0 7°, 00 Appendix 11. (Cont'd) Appendix 11. (Cont'd) s 0.44 OOF 05- » 1 1 \ Dl - -169*. * 0 .2143F -01* 02 0 .35 OOF 05-FRATloiCOE rF.). 53.51 FPO-jmcoccc, |- 0>-> | < STO , ?.\3 r., r Civ5 T< = ST'l,F=-p ,cor r t=„ = 0.3043E-02 5Tn.E5o, n; = 4-->3„ •0.26 0OE 1 2 05- < OI'FMIV-WATSON STA." 0,3197 . . A'.iriicoc-RFIAT I'lN Cn"=r. . 0. 5742 ! - 1 0.1700E 05-1 1 - I 1 t 1 1 • 3030, 1 1 1 1 . 1 1 7 1 1 * 1 X I 1 72 ,41 ) i I 1 12, 311 1 1 2121 ? i -1000-1 13213*2577)1 7 1 )  — 9 -O.2O0OE 05 0.4600E 06 0 . 0,2700E 06 0.7000F 06 9400E 06 0 , H . 3 0 C 07 0 4 4 OOF 05- 1 1 1 0! - -177*. » 0.747? = -01 » 03 0.3500F 05-PRATfMCOECF. 1' 33.64 F ^ i n i f n c r f O.fiO.TuJ ! < STO.FR3.CONST,* 1335. S T n . F S ' . C i f ^ F . ' 0.3T76F-0? STO.coc). ni 6r'71. 0 . ?< OOE 05- < K ? 0 • 0." 2 11 O'.PIIN-WITSON 5TA. ' 0. 3037 AIITOCORS ev ,<T ION C.OVF. -> 0.5306 | . 1 0, 1 7 OOF 05- . * 1 * 1 * 1 1 1 , 8000, 1 Ir 1 I . 1 1 1 1 2, 1 11 1 2 3. 7171 1 1 1 1 2 . 1 3 71 3 131 1 -1000. f ] 332,1 *2527 • 111 1 J.J 1 1 -0.2000E 05 0^4000^ 06 0 . 0 . J900E 06 0.6100E 06 8?00? 06 0 . 1030E 07 Appendix 11. (Cont'd) 0.4400F 05- 1 I > — n i . -ms-H. • 1506. * 04 0,3500F 05-FRATlotCOEFF. )= 21,76 | < < STO, F 3 7 .CO'IS T, = 1706. ST0.P5R.CfFfe , . 122.8 STO.E"?. 0' " 7736, 0.26OOE 05- 2 RSI = 0 . 7 1 1 8 Oi|»1i\'-'.JATSP\' STA., = 0, 8567. ."V'JKCLRILATIO-: CO-=n. . .0.57 07 j ! 1 0.1700F 05-1 1 1 l 8000, ! l 1 i j 1 3 .2 7 3 . 3 1 2 I 3 ? 1 5 7 7 1 -1000. 1 7 - > ,7 7 2 2 2 2 —£ 1. 000 2 . 600 4, 200 5, POO 7, 400 9, 000 . i ! • • ~ ~ i ro UJ Append i x 11. (Cont'd) ni « '421. » -135„Q * 03 F R U I O ( c n r C F . i« o.9?95F-o?. F p R 0 3 ( C O P C F > ) » 0 .3351  S T O . F ' R R . C O N S T , ' 2175. S T O . F R " . C O P F F , . 1410. ST^,ERP. 0' * 73 2 5,  RSO ' 0, 330? OM^OIN-WATSON STA, = 0, 3864 A'ITQi".2PRFlAT?TJ COF-F, s 3,5562 HI = 7739. 4- -?53.1 • 08 FRAT!0(C0FFr,|. 1,0'0 -FPO,nP,(CnF--F.)» 0.3167  STD.F.RR.CONST,» 2531. STO.Fp.p..COFFF. * 25?.5 •STQ.FRP. 01 . 73 T 6 . PSO ' 0.01 24 OIJP'HM-WATSON STA. . 0. 3993 .AljTnCOPP FJ AT_j_ON COT ~ p , * 0,54 34 01 - 7"54c ^ -1733. * 04 FRATIOtCOFFF,)» 13.70 F P 3,0 3 K' l e J iF U . * Qjji 017 STO.ERR,CONST,' 11 67. STO.^Rs.COrEE. ' 520.7 <rn .H ' . 01 - ",4 0.  PSO ' 0. L! 6" OIFRIN-HATS'ON S T A , ' 0,7643 . AIJ r or o R R EL A T J 0 \i _C0" c :. 01.6 !_63__ 01 » 0.1 776 ! : 05* -323, 4 * 03 FRATIOtCOEFF. )« 4,046 _ FPRnqtCQf 0.0431  STD.ERR.CONST.' 3333. STO.ER°.CnFFF,« l<i6,R S T O , - P P . Dl » 77 34.  RSO = 3,3474 DURRT'l-'WATSON STA, * 0, 9T10 A'JTnroppPl ATT ON C O ^ F . » 0.3330 01 » 3704, «• 1346. * 07 FRATIOtCOEFF.)' 0. 4501 E-01 FfoO'llCOEF". I ' 0,315)  STO.FRR.CONST,' 72 2 7. STO.pop.COEPF.' 3*00, . S T o.rR", 01 * 7T.3.  RSO ' 0,0005 . 0II"1IN-WA TSDN ST4„' 0,3380 _ n'.ITOrOF PEL AT !0N C O " F , _ « P ^ V S 54 01 u 5956. » -17.0? * Ql 0 FDAT10 I COEFF. 1» 0.1744 FPR03(C0EFF.)* 0.7732 STO.TPR.CONST, « 27 3"*. STO.EP.R.COFFF.* 43, 27 STO-.TP". r>\ 2 7". °i 0S0 » 3,3 115 0UR3 IN-WATSON S T A , ' 0.8884 j i i i T n r n p p r i A T T O N r.O^-c. . 0. 5534 Offifou^ HIM ft 6- tieA/TJ op -rue Cut A re M &>~c. 0, tv / tfugrfuM t/e-ru. ~JT, Sera /fe_\ <{-C0 : •Zr?~3l. Asso c; &-rey> s-r/euc-ro seers WITHIN 'rue i^^itje: cia^Ye •1i¥ (s-z-aJ-SiJ-Sii. /^ m> / T J IN Co MPL,£-~re vTy«/2/OA/-a : # / € e v / ^ « v Meow** T. ftstt. r€es. 

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