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The effect of coloured lights on the reproduction of Wistar rats Aldous, John Gray 1941

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LE .3 & 1 THE EEEECT OE COLORED LIGHTS Off THE REPRODUCTION 03F WIS TAR RATS 37 John Gray Aldoua A Tlieala submitted; i n p a r t i a l f u l f i l l m e n t f o r the degree of MASTER OF ARTS Department of Biology and Botany, The U n i v e r s i t y of B r i t i s h (JoiumMa, August, 1941 Table of Contents Page AciaiQwledigefflerits I n t r o d u c t i o n x Review of l i t e r a t u r e — . — — 3 ; "Experimental — . — — — 15 R e s u l t s . . — 19 Diacusrsion and Conclusion — —- 2& Summary ~~ • - — — — — • — • - 3 0 B i b l i o g r a p h y • — — - — ~ . — — 31 Graphs --•— •* -— 3 ^ Acknowledgements The author wishes to> express h i s a i n c e r e s t thanks to the f o l l o w i n g persons r To Dr. JV A l l a r d y c e , Department of Biology amd Botany, under whose s u p e r v i s i o n t h i s i n v e s t i g a t i o n was c a r r i e d out, f o r h i s c o n s u l t a t i o n s , valuable advice and as s i s t a n c e , and e s p e c i a l l y f o r performing the a c t i v i t y t e s t s on the r a t s during the summer of 194I. To Dr. A;.K. Hutchinson, Department of Biology and Botany, whose u n t i r i n g i n t e r e s t and assistance i n obta i n i n g apparatus made the p r a c t i c a l and the t h e o r e t i c a l side of the i r r a d i a t i o n work much e a s i e r . To> Dr. A..H. Croaker, Department of Phy s i c s , f o r h i s "many hours spent incchecking the spectroscope, galvanometers, and thermocouples, f o r h i s help i n determining transmission spectra and g e n e r a l l y b r i n g i n g the t e c h n i c a l side of r a d i a t i o n w i t h i n the scope of a physiology student. To Miss. J.M. P r a t t , who d i d a l l the feeding and cleaning of the cages during the season 1940-41. To Mr. B i l l Cooper, who c a r r i e d on the experiment during the summer of 1941, and without whose help t h i s work would be incomplete. To these and many others, both v/ithin and outside the Department of Biology and Botany, the author owes much; t h e i r i n t e r e s t and enthusiasm during the two years of the i n v e s t i g a t i o n have made the work very worth while. THE EFFECT OF COLORED LIGHTS UPOH THE REPRODUCTION OF WISTAR RATS The f a c t that l i g h t , monochromatic or heterochromat-i c plays an important part i n the l i f e processes of organisms, has been r e a l i s e d f o r w e l l over a century, but the discovery of i t s r o l e i n c o n t r o l l i n g organic development and reproductive cycles both i n pl a n t s and animals has only been f u l l y r e a l i z e d during the l a s t decade of s c i e n t i f i c h i s t o r y * The c o n t r o l of times of maturity, e s p e c i a l l y i n p l a n t s , has had many a p r a c t i c a l a p p l i c a t i o n s i n c e , by i n c r e a s -ing: or decreasing the amounts of d a y l i g h t , plants may be made to bloom any d e s i r e d time of the year. Such treatments have a l s o been found e f f e c t i v e i n i n c r e a s i n g the development of root s i n tuberous crops, or leaves i n l e a f y crops; moreover, both types of development may be enhanced by g i v i n g a pre-treatment of the r e q u i r e d l e n g t h of day, fo l l o w e d by a r e t u r n to normal. This "carry-over* e f f e c t has been put to much use i n seed v e r n a l i z a t i o n , whereby, besides the above mentioned r e s u l t s , r e s i s t a n c e to f r o s t i n j u r y i s increased. S i m i l a r p h y s i o l o g i c a l e f f e c t s have been noted i n regard to animals, but although a p r a c t i c a l a p p l i c a t i o n i s perhaps not so evident, except f o r such cases as i n c r e a s i n g egg production from hens by using supplementary a r t i f i c i a l l i g h t , negertheless, human beings aye exposing themselves to a l l kinds of r a d i a t i o n s , e s p e c i a l l y since the development of a r t i f i c i a l l i g h t i n g ; therefore i t i s of utmost importance that t h i s subject be studied i n i t s minutest d e t a i l * I t s p h y s i o l o g i c a l importance l i e s not only i n i t s method f o r c o n t r o l of animal breeding, but f o r those e f f e c t s upon human physiology of which we are yet unaware» The s e l e c t i o n of the wavelengths of r a d i a t i o n responsible f o r these organic changes, has provoked much controversy, and i t i s w i t h the purpose of adding to the already accumulated data that t h i s work was undertaken. Review of l i t e r a t u r e As might be expected, experiments that have had a d i r e c t a p p l i c a t i o n to commerce have received by f a r the greatest a t t e n t i o n * Thus, much of the work done on the p h y s i o l -o g i c a l e f f e c t s of r a d i a t i o n has f a l l e n under the heading of photoperiodic response* L i t t l e reference w i l l be made h e r i n to such responses i n p l a n t s , f o r the l i t e r a t u r e upon the subject i s voluminous, and only a s m a l l p o r t i o n of i t bears any r e l a t i o n to t h i s work. For the same reasons the e f f e c t s of u l t r a v i o l e t l i g h t w i l l only be noted i n passing; e x c e l l e n t papers and b i b l i o g r a p h i e s may be found i n Duggar (13) and Heyroth (20). The importance of i n c i d e n t r a d i a t i o n i n c o n t r o l l i n g organic development was studied i n d e t a i l by Brown, Pearce, and Yan A l l e n ( 4 ) . They noted a seasonal f l u c t u a t i o n of organ -weights i n r a b b i t s , which remained constant over a p e r i o d of s e v e r a l years* Following t h i s , the results, of post-mortem and c l i n i c a l observations were reported by Pearce and Yan A l l e n ( 3 4 ) * The r a b b i t s were maintained i n three environments, constant i l l u m i n a t i o n , s o l a r i l l u m i n a t i o n and darkness. Those l i v i n g under constant l i g h t c o n d i t i o n s , tended to have smaller r e l a t i v e weights than the c o n t r o l s , p a r t i c u l a r l y n o t i c e a b l e i n the l i v e r , kidneys, g a s t r o i n t e s t i n a l mass, t h y r o i d , para-t h y r o i d s , hypophysis, p i n e a l body, spleen and rep r e s e n t a t i v e lymph nodes* The other organs, i n c l u d i n g the h e a r t , b r a i n and t e s t i c l e s , showed a s i m i l a r trend, but l e s s marked. The thymus proved to be a consistent exception} f o r con s t a n t l y •7 i l l u m i n a t e d animals the weights were higher and f o r "dark" animals the weights were lower than the c o n t r o l s , A second general e f f e c t was noted, i n that constant i l l u m i n a t i o n tended to make f o r greater s t a b i l i t y of organ weight, i.e», the c h a r a c t e r i s t i c f l u c t u a t i o n of the c o n t r o l s was absent. This e f f e c t was evident to a considerably smaller degree i n the "dark"' animals. Brown (2,3), g i v e s the r e s u l t s of a s i m i l a r exper-iment wherein three groups of r a b b i t s (white, black, and dark colored) were subjected to three d i f f e r e n t l i g h t c o n d i t i o n s , namely, c o n t r o l ( d a y l i g h t ) * darkness, and constant i l l u m i n a t i o n from twenty standard crown g l a s s neon tubes, 6 f t . , i n le n g t h , 5/8 i n , diameter, emitting the greatest p o r t i o n of r a d i a t i o n i n th© region 58000-7600 A., w i t h a small short-wave emission at -'3370-3620, concentrated p r i n c i p a l l y a t 3460-3430 A., The general trend of organic development i n these animals was much the same as was: reported by Pearce and Van A l l e n , but, perhaps more s i g n i f i c a n t i s the observation t h a t , f o r some organs, there i s unquestionably a d i f f e r e n c e i n the weight f o r the "- l i g h t " and "dark" animals, when compared to the co n t r o l s 5 those i n the dark showed greater values than those i n the l i g h t . S i g n i f i c a n t a l s o i s the f a c t t h a t , whereas the white animals were below normal, the b l a c k animals wei?e above normal. Outstanding i n the analyses of organ weights are the r e s u l t s from the t h y r o i d . For the c o n t r o l animals i t was found that there e x i s t e d a d i s t i n c t d i f f e r e n c e i n the weights of t h i s -5. organ from- the white, and the black or dark colored animals. This r e l a t i o n a l s o h e l d f o r the " l i g h t " and adark* d i v i s i o n s of Groups I (white) and I I (black) hut not f o r Group I I I (dark c o l o r e d ) . Sheard and Higgins ( 3 7 ) w h i l e studying the e f f e c t s of supplementary diets; w i t h Zf& cod l i v e r o i l , f e d to c h i c k s under various l i g h t f i l t e r s , r e p o r t that h y p e r p l a s i a of the para-th y r o i d s i n v a r i a b l y occurred under blue and amber g l a s s . They suggest that t h i s overdevelopment takes place i n the i n t e r e s t of maintaining normal growth; u l t i m a t e l y , however, t h i s i s not adequate and subnormal weight f o l l o w e d by signs of r i c k e t s , occurs. In a l a t e r paper (38) the same authors confirm the a b o v e r e s u l t s , f i n d i n g that the p a r a t h y r o i d enlargement was due to h y p e r p l a s i a rather than hypertrophy. Moreover, there was? no organic i n d i c a t i o n , e i t h e r i n blood calcium, or blood phosphorus, or general a c t i v i t y , that such h y p e r p l a s i a had developed. Both the longer and the s h o r t e r wavelengths of s u n l i g h t were found e s s e n t i a l f o r normal p a r a t h y r o i d develop-ment. Somewhat c o n t r a d i c t o r y to t h i s , are the observations of 3? a i r h a l l (1.5), who found that r a t s , on a normal calcium d i e t showed a higher calcium content when i r r a d i a t e d w i t h u l t r a - v i o l e t l i g h t . Both on a low and normal calcium d i e t , the r a t s showed a higher degree of u t i l i z a t i o n of t h i s element under i r r a d i a t i o n . ' Following extensive work on the e f f e c t of l i g h t on the sexual c y c l e of b i r d s , Bissonnette (5) came to the co n c l u s i o n that "progressive t e s t i s changes i n s p r i n g , may be helped by the r e l a t i v e strength of red rays i n the s u n l i g h t of that season, and the r e g r e s s i o n beginning before June 21 i n nature may be due to the r e l a t i v e excess of i n h i b i t o r y or l e t h a l green (and perhaps vio.iet and u l t r a v i o l e t ) rays i n summer and e a r l y autumn." In t h i s study, Bissonnette found that birds, under r e d l i g h t 2.6 f.c.) reached complete spermatogen-e s i s but no white or green l i g h t e d ones d i d . Working w i t h female f e r r e t s , M arshall and. Bowden (28) report that heat rays and near i n f r a red are comparatively i n e f f e c t i v e i n a c c e l e r a t i n g the recurrence of oestrus j. the e f f e c t , beginning w i t h the red i r r a d i a t i o n (6500 A) and extend-i n g throughout the v i s i b l e to the near u l t r a v i o l e t (3600 A j , appears to depend more on i n t e n s i t y than wavelength. H i l l and Barkes (21, 22) d e f i n i t e l y e s t a b l i s h e d the f a c t that the response o f female f e r r e t s i n anoeatrus, to a d d i t i o n a l i l l u m i n a t i o n , was dependent upon the presence of the a n t e r i o r p i t u i t a r y . Moreover, (Browman ( I ) ) s t i m u l a t i o n of t h i s organ i n a l b i n o r a t s was shown to depend upon the presence of the r e t i n a , as animals w i t h r e t i n a s removed, or o p t i c nerves severed, f a i l e d to respond to increased a r t i f i c i a l i l l u m i n a t i o n (Browman ( I ) ) . Recently, M a r s h a l l ' s (2?) r e s u l t s on u l t r a v i o l e t i r r a d i a t i o n p o i n t out the p o s s i b i l i t y that such i r r a d i a t i o n may cause a r e l e a s e of a f o l l i c l e - s t i m u l a t i n g hormone, as w e l l as a l u t e i n i z i n g hormone* from the p i t u i t a r y , the former always being dominant* Considering that f e r r e t s o n l y ovulate a f t e r c o i t i o n , "-the existence of w e l l defined l u t e a l t i s s u e i n the ovary of the f e r r e t which had never been w i t h a male, may probably have been due to the release of the l u t e i n i z i n g hormone consequent upon u l t r a v i o l e t i r r a d i a t i o n . " Thus, the course and probably the consequence of ra d i a n t energy s t i m u l a t i o n on reproductive organs was f a i r l y c l e a r . H a r r i s (18} found that d i r e c t e l e c t r i c a l s t i m u l a t i o n of the p i t u i t a r y i n female r a b b i t s , induced o v u l a t i o n 15-40 hours l a t e r , and sometimes r e s u l t e d i n the formation of c y s t i c and haemorrhagic f o l l i c l e s * S t i m u l a t i o n of the hypothalmus gave s i m i l a r r e s u l t s . Matthews (29), co-nducted an i n t e r e s t i n g experiment by which i t was shown that o v u l a t i o n i n pigeons was a d i r e c t response to a v i s u a l s t i m u l u s . B i r d s were set up i n the f o l l o w i n g s e r i e s * Double cages Ho* I Male and female w i t h no p a r t i t i o n ( c o n t r o l ) • Ho. Z Male and female w i t h glass p a r t i t i o n . No. 3. Two females w i t h g l a s s p a r t i t i o n . Bo. 4 Two females w i t h no g l a s s p a r t i t i o n . S i n g l e cages Ka. 5 Female alone w i t h m i r r o r . -8-' No* 6 Male alone w i t h m i r r o r * No. 7 Female alone without m i r r o r * A l l the females* except Ho* 7 ovulated normally? her "subsequent o v u l a t i o n * when placed w i t h a male showed her to be a normal b i r d * The response of the female i n cage No. 2 showed such o v u l a t i o n was not a t a c t i l e response* while the response of No* 5 confirmed the expectation that o v u l a t i o n was v i s u a l l y s t i m u l a t e d . V i s i b l e r a d i a t i o n has been found, by Luce-Glausen and Brown (25) to promote growth i n r a t s ; i n f r a r e d e f f e c t s were d e f i n i t e but l e s s i n degree. Rats confined to dark con d i t i o n s stored Vitamin A more r e a d i l y than those exposed to i r r a d i a t i o n * I t i s suggested that r e t i n a l s t i m u l a t i o n may pl a y a part here* Continuing the work i n r e l a t i o n to i t s e f f e c t upon reproduction, the same authors (26) found that v a g i n a l opening and the onset of oestrus were delayed when r a t s were confined to darknessj those i r r a d i a t e d w i t h the v i s i b l e spectrum matured e a r l i e r than those under normal experimental co n d i t i o n s S u r v i v a l of young was higher under v i s i b l e r a d i a t i o n although energy equivalents were one t h i r d of those f o r the near i n f r a r ed, where the above e f f e c t s were intermediate between those of v i s i b l e and f a r i n f r a red* P e r i o d i c f l u c t u a t i o n s i n the t e s t i c l e s and proatas.tes of r a t s has been noted over a p e r i o d of four years by D e l G a a t i i l o and Pinto (10). Decrease i n t e s t i c l e weight occurred i n the autumn, c l o s e l y followed by the p r o s t a t e ; during: the winter the t e s t i c l e recovered, w h i l e the seminal v e s i c l e s and prostate increased g r a d u a l l y , reaching a peaft during the summer. Heat (36C) and c o l d (I6G) and l i g h t of d i f f e r e n t i n t e n s i t i e s (50 and 400 c.p.) caused l i t t l e v a r i a t i o n frominthe c o n t r o l s . Bisaonnette (6) found that male blue jays responded to a d d i t i o n a l i l l u m i n a t i o n i n December by almost complete spermatogenesis and a c t i v a t i o n of epididymes to complete breeding c o n d i t i o n i n 28 days. D e f i n i t e e f f e c t s may thus be a t t r i b u t e d to i n c i d e n t r a d i a t i o n , although as yet i t i s a matter of conjecture whether these e f f e c t s are due to wavelength, i n t e n s i t y , or increase "in photoperiod. Colored l i g h t s are known to have an e f f e c t upon human behaviour, but l i t t l e i s known through what paths these e f f e c t s manifest themselves, or whether constant exposure (through such media as neon l i g h t s , or room c o l o r i n g s ) to such conditions i n s t i g a t e harmful organic changes. The t h e o r i e s of the b i o l o g i c a l e f f e c t s of r a d i a t i o n have been very improperly developed, due, i n the main to inadequate experimental s u b s t a n t i a t i o n . A few, however, are worthy of note. Deasauer ( I I ) , working on the s u p p o s i t i o n that strong b i o l o g i c a l e f f e c t s of r a d i a t i o n could not be due to uniform thermal increase i n the m a t e r i a l , but r a t h e r to -10-d i s c r e t e points of heat f o c i , substantiated t h i s view i n regard to albumin and pseudoglobin. He found that a f t e r i r r a d -i a t i o n , these p r o t e i n s were coagulated and became v i s i b l e I n Brownian movement, when examined under the ultramicroscope. Many workers have hi n t e d that I n j u r i o u s e f f e c t s may be a s c r i b e d to the d e s t r u c t i o n of enzymes| but here a d i s c r e p -ancy a r i s e s , . C o l l i e r and Wasteneya (8) report that u l t r a v i o l e t r a d i a t i o n (2000-3130 A.) destroys impure urease, malt amylase, pepsin and plasma phosphatase. Urease i s a l s o destroyed when exposed to wavelengths of 750Q-I4,0Q0A.. V i s i b l e r a d i a t i o n stimulates s t a r c h amylase d i g e s t i o n , while pepsin-ovalbumin d i g e s t i o n proceeds r a p i d l y , although i t has no e f f e c t on the enzymes alone. Accordingly i t i s suggested that the c o l l o i d a l s u b s t r a t e absorbs the r a d i a t i o n , the energy of which i s then degraded i n t o k i n e t i c form, w i t h consequent increase i n the r a t e of r e a c t i o n . In d i r e c t contrast to t h i s , P u l l e r (16) has shown that enzyme a c t i v i t y a c t u a l l y increases, i n t i s s u e s which have been i n j u r e d by d e s t r u c t i v e wavelengths. Thus i t i s demonstrat-ed that the i n j u r i o u s e f f e c t s of u l t r a v i o l e t must be traceable to some p h y s i o l o g i c a l dearrangement other than i n a c t i v a t i o n of enzymes. I t was also shown that any i n a c t i v a t i o n as r e s -u l t e d from i r r a d i a t i o n was due, i n considerable degree, to the i n f r a r e d . Hurnberger (32) a t t a c k s the problem from an Ion-I z a t l o n point of view, and suggests t h a i a l l r a d i o - b i o l o g i c a l r e a c t i o n s f o l l o w a common p a t t e r n , v i z . , the energy i s absorbed and d i s s i p a t e d by the decomposition of water i n t o negative and p o s i t i v e i o n s . This energy i s t r a n s f e r r e d to the c e l l c o n s t i t -uents, where physico-chemical r e a c t i o n s are i n s t i g a t e d , r e s u l t -i n g i n de v i a t i o n s from the normal physiology of the c e l l . Whether " s e l e c t i v e effects'* are j u s t apparent, or whether they are r e a l and how much they depend upon the primary i o n i z a t i o n or the secondary energy exchange, are questions which the author does not attempt to answer, Packard 133) i s of the opinion that x-rays and °<-rays do not d i r e c t l y s t i m u l a t e the normal a c t i v i t i e s of the c e l l . The primary e f f e c t i s always an i n j u r y from which the c e l l may recover p e r f e c t l y , the a c c e l e r a t i o n of some normal - c e l l processes, which has been observed, being a tjasaporary response to i n j u r y . I t i s , however, p o s s i b l e that the products of c e l l i n j u r y , whether u l t r a v i o l e t or x-rays are used, may contain growth promoting substances which have the power of s t i m u l a t i n g uninjured c e l l s . I t i s w e l l known that i n j u r y to pla n t t i s s u e r e s u l t s i n an immediate r i s e i n that t i s s u e s ' r e s p i r a t i o n and, i n the l i g h t of Packard * s suggestion, i t may w e l l be that the st i m u l a t o r y e f f e c t s of i r r a d i a t i o n are o n l y secondary, the primary r e a c t i o n being i n j u r y , f o llowed by an increase i n the r e s p i r a t o r y process and a subsequent increase i n c e l l u l a r a c t i v i t y due to s t i m u l a t o r y substances secreted. -12-From a study of the absorption spectra, "both of b i o l o g i c a l m a t e r i a l s and t h e i r chemical c o n s t i t u e n t s , s p e c t r a l regions, where a p h y s i o l o g i c a l r e a c t i o n to i r r a d i a t i o n might be expected, soon become apparent, because i n these regions, there i s a high absorption of the i n c i d e n t r a d i a n t energy. Proteins and amino acids form an important part i n the s t r u c t u r e of p h y s i o l o g i c a l processes. Consequently, i n those regions, where such substances absorb s t r o n g l y , one would expect strong r e a c t i o n s to i r r a d i a t i o n . Such a c o r r e l a t i o n manifests i t s e l f i n many ways. A l l p r o t e i n s show a prominent absorption i n the reg i o n 2650 A., whil e t h e i r c o n s t i t u e n t amino acid s ( i n p a r t ) have been found to account f o r t h i s band. The phenyl alanine band spreads from 2500 to 2700 A.? t r y o s i n e shows a maximum a;t 2840 A., tryptophane a t 2920 A., while such others as asparagine, h i s t i d i n e and a r g i n i n e show bands covering t h i s area. The m a j o r i t y tend toward a minimum at 3000 A. (24}. Turning now to the e f f e c t s of r a d i a t i o n i n t h i s r e g i o n , we f i n d that b a c t e r i c i d a l a c t i o n against S. raureus occurs over 2250-3130 A., w i t h a maximum a t 2600-2700 (17). P r o t e i n coagulation i s highest: a t 2600 A.; v i r u s suspensions are i n a c t i v a t e d a t 2650 A (35); erythema production shows two maxima, one a t 2500, the other a t 2950, w i t h minima a t 2750 and 3I0Q A. ( 7 ) , hemolysis of blood c e l l s occurs a t 2450, w h i l e the wavelengths l e t h a l to _S» marcescens and A s p e r g i l l u s niger are l o c a t e d a t 2650 and 2537 A. r e s p e c t -- r s -i v e l y (40). i Maximum l e t h a l e f f e c t i v e n e s s to Q h l o r e l l a v u l g a r i s was found a t 2650 and 2800 A. (31'). while mutations i n Trichophyton mentagrophytea have been induced by i r r a d i a t i n g w i t h the band 2537-2650 A» (1.4). Spores of U s t i l a g o zeae show a maximum s e n s i t i v i t y at 2652, and a minimum (no k i l l i n g ) at 3130 A. (23). ghaetomiuiB, Drosophila, paramoecium and B. c o l i , a l l t reated under the same conditions show a maximum s e n s i t i v -i t y a t 2652, and a minimum (no k i l l i n g ) at 3130 A. (23). On the other hand, the r e g i o n of greatest spore production f o r Macrosporiurn f u s a r i u m i l a y i n the wavelength i n t e r v a l of 2535-2800 A.j s l i g h t s t i m u l a t i o n occurred a t 3120, while no e f f e c t was noted from 3334 A., on (36). Aside from t h i s l a t t e r case, i t may g e n e r a l l y be concluded that the r e g i o n 2500-2800 A., i s l e t h a l to most organisms, p o s s i b l y i n view of the f a c t that strong p r o t e i n absorption occurs here. Such d e f i n i t e conclusions regarding the region l y i n g around 3000 A., cannot be drawn. Detwi l e r (12) reported t h a t i r r a d i a t i o n embracing 2700-3200 A. caused delay i n germination and pronounced s t u n t i n g of seedlings of Ribes r o t u n d i f o l i u m . Withrow and Benedict showed that the removal of the 2900-3100 band was detrimental to the growth of tomato and Goleus p l a n t s . They intimate that t h i s might be a growth promoting r e g i o n (39). -14-Perhaps, t h i s discrepancy i s not so hard to overcome, i f one r e f e r s to the region inducing the production o f erythema; maximum production occurs at 2950, and minimum at 3050 A., In other words a range of 100 A. at 3000, may determine a l l or none. Detwiler used a very wide range of wavelengths (2700-3200) and p o s s i b l y the s t i m u l a t i n g e f f e c t that Withrow and Benedict reported was overcome by the l e t h a l e f f e c t s of the shorter wavelengths..uThis argument a p p l i e s to m much of the work done i n the u l t r a v i o l e t , i . e . , the wavelength i n t e r v a l s have not been s u f f i c i e n t l y narrow tompermit i n t e r -p r e t a t i o n of i n d i v i d u a l bands to be made. Experimental This study of the i n f l u e n c e of c o l o r e d l i g h t s on the development of r a t a was f i r s t undertaken i n the P a l l of 1959. At that time* a p a i r of r a t s , selected from the remain-der of Miss. B a l e 1 s stock were placed a t s i x weeks of age i n each of eight cages. These cages were constructed as f o l l o w s J Sides and back of galvanized i r o n , w i t h f r o n t , top, and bottom of 3/8 i n . mesh galvanized wire netting? the hinged f r o n t was f i t t e d w i t h galvanized i r o n clamps f o r holding the i n v e r t e d Erlenmeyer d r i n k i n g f l a s k s . O v e r a l l dimension: 12 i n . long, 10 i n . wide and 8 i n . deep. The whole cage f i t t e d i n t o a galvanized i r o n pan, -| i n . deep, i n which was placed a double thickness of paper toweling, covered w i t h sawdust, leaning of these cages was done once a week. The f i l t e r s c o n s isted of sheets of colored cellophane stretched over wire frames and placed d i r e c t l y i n f r o n t o f the cage. On a l l cages s i n g l e sheets were used, but on the black about f o u r sheets were found necessary to exclude the l i g h t . Two determinations of the s p e c t r a l c h a r a c t e r i s t i c s of these f i l t e r s were madej t h e i r absorption spectra showed the f o l l o w i n g regions of transmission? -0 This stock was used f o r two years i n thymus feeding experiments. - r e -V i o l e t . 5800-6800 A., peak at 6400 A., s l i g h t t r a n s -mission at 43iOQ-4700 A. Blue 4900-6500 A.» peak at 5400 A., s l i g h t t r a n s -mission at 4100-4700 A* Green 5000-6500, peak at 5300 A. Yellow "J 5200-6500 A., broad peak at 5500-6500 A., Grange J orange transmits s l i g h t l y more red Red 5800-7000 A., peak a t 6200 A. These measurements were made on a H i l g e r Quartz, spectrograph using: Wratten Wainwright Panchromatic p l a t e s ; the l i g h t source was a 60 watt Mazda f r o s t e d globe, one f o o t from the c o l l i m a t o r s l i t . The 100 W. Mazda bulbs used i n the i r r a d i a t i o n of the r a t s , would of course emit more of the shorter wavelengths, but the c u t - o f f s of the cellophane f i l t e r s would l i e approximately i n the same re g i o n s . Because c h a r a c t e r i s t i c s of the Wratten Wainwright p l a t e s used, no wavelengths longer than 7000 A. could be recorded, so t h a t , although the red, orange and yellow f i l t e r s showed c u t - o f f s a t 7000 A., i t does not f a l l o w that near, or f a r i n f r a r ed was excluded by these f i l t e r s . Clear cellophane transmits 50% of the i n c i d e n t i n f r a r ed from 10,000 A. to 24,000 A. (Cartwright 19), so that there i s a strong p o s s i b i l i t y that the r a t s were r e c e i v i n g much more of t h i s part of the spectrum than i s shown i n the photographic p l a t e s . Transmission of the i n c i d e n t r a d i a t i o n was -17-measured w i t h a H l l g e r thermocouple, and galvanometerj the r e s u l t s agreed w e l l w i t h those found i n the l i t e r a t u r e . Beyond 3000 A. there i s a l i t t l e change i n transmission, and even as low as 2-000 A.., $6% of the i n c i d e n t r a d i a t i o n i s transmitted (Phys. Rev. S i r I I I 4 : 1928). L i g h t from-two 100 watt. Mazda lamps, at a distance of 10 f t . , formed the source of i l l u m i n a t i o n . A temperature of 70 E. was maintained "by two thermostats* one c o n t r o l l i n g an e l e c t r i c heater, the other, a f a n f o r a i r c i r c u l a t i o n . In the F a l l of 1940 new f i l t e r s were set up i n f r o n t of the cages, and an. automatic time s w i t c h was. i n s t a l l e d to ensure uniform periods of 10 h r s . i l l u m i n a t i o n , commencing at 8J00 a.m. and terminating a t 6 sQQ ,p.m.. Since post morterns had "been performed on the r a t s from the previous season, a breeding stock of Wis-tar a l b i n o r a t s was obtained from the Department of B a c t e r i o l o g y , and from these l i t t e r s , two or three females plus one male were placed i n each of the experimental cages at 3 weeks of age. Feeding was c a r r i e d through 6 days per week, each r a t r e c e i v i n g three P u r i n a Ghow Checkers per day, except during the period of r a i s i n g a l i t t e r , when the female was fe d f i v e per day. This d i e t was o c c a s i o n a l l y supplemented by green l e a f y vegetables and c a r r o t s . At the time of c a s t i n g of l i t t e r s , a l l other r a t s were removed to a new cage w i t h a new f i l t e r , l e a v i n g the mother alone, w i t h the l i t t e r , u n t i l reaching three weeks of age, a t which time, the female was returned to the male. At t h i s time, one male and two females were select e d from the l i t t e r and placed i n a new cage w i t h a new f i l t e r . In t h i s way the r a t s were r a i s e d , generation to generation under the same wavelengths of i r r a d i a t i o n . The r a t s were weighed once a week u n t i l 70 days of age, t h e r e a f t e r the females were weighed twice a week i n order to o b t a i n a peak weight previous to l i t t e r c a s t i n g . The l i t t e r s were handled as l i t t l e as p o s s i b l e , therefore weighing, i n toto was performed only at b i r t h and at three weeks of age. The number of deaths, e i t h e r from m a l n u t r i t i o n or from cannibalism were noted f o r each l i t t e r . Resulta General As was expected from a review of the r e s a l t s i n the l i t e r a t u r e , l i t t l e d i f f e r e n c e was noted between the weights of the r a t s r a i s e d under the d i f f e r e n t f i l t e r s , although those r a t s that came from a small l i t t e r u s u a l l y had an advantage i n weight over those which came from a l a r g e one; the advantage, however, became n e g l i g i b l e a f t e r an age of 80 days or so (Tables I , 4 ) . Averaging a l l the l i t t e r s under i r r a d i a t i o n , together, there appeared to be some r e l a t i o n s h i p between i n d i v i d u a l weight a t b i r t h and numbed i n l i t t e r , but t h i s was not n e a r l y as constant as was the r e l a t i o n between i n d i v i d u a l three week weight and number s u r v i v i n g to three "weeks. The l a t t e r values when p l o t t e d against each other (see f i g . I ) showed a s t r a i g h t l i n e r e l a t i o n s h i p between l i t t e r s of 3 and 6; below 3 the i n d i v i d u a l weight went up s l i g h t l y , above 6 the r e l a t i o n s h i p assumed a curve, w i t h an apparent minimum of 18 g r . per i n d i v i d u a l . When weights were a as low as t h i s , i t $aa o f t e n found impossible to wean the young at three weeks, so u s u a l l y the mother was l e f t i n the cage u n t i l such time as the young appeared able to feed themselves. C r a z i e r (9) found that the r e l a t i o n between b i r t h weight and l i t t e r s i z e i n mice could be expressed as Wt» alf^ where, f o r homogeneous data, k has the n o n s p e c i f i c value of 0,83, No such r e l a t i o n s h i p could, be found, i n the present r e s u l t s ; v a r i a t i o n i n the weights f o r l i t t e r s of the same s i z e may have been i n f l u e n c e d to some extent by the f i l t e r e d r a d i a t i o n . A c t i v i t y t e s t s were performed on a l l P 0 generation r a t s by dropping them i n t o a rubber diaphragm; movements were recorded on a smoked drum r e v o l v i n g a t a slow speed, so that the tracing- represented about 90 seconds d u r a t i o n . These are reproduced i n P i g , 2 f o r the males only, as an u n c o n t r o l l a b l e and indeterminate e r r o r was introduced i n the case of the females, whose d i f f e r e n t stages i n the\oestral c y c l e provided wide d i f f e r e n c e s w i t h i n the same c o l o r group. Generally speaking, the males were more a c t i v e than the females. Greater a c t i v i t y was found i n the Black "and Red males, although the former observation was a l i t t l e s u r p r i s i n g i n that t h i s r a t appeared q u i t e normal when handled. White and Orange proved to be the l e a s t a t t r a c t i v e , f o l l o w e d c l o s e l y by the Blue, Yellow, V i o l e t and Green. Fp Reproduction (Table 2) In general the e f f e c t s of the i r r a d i a t i o n on the d i f f e r e n t phases of reproduction appeared to bear some r e l a t i o n to i n c i d e n t wavelength^ Taking the Black as the c o n t r o l animals, White l i g h t i n h i b i t e d production o f \ l l t t e r s to the extent that a t the end of 212. days only one l i t t e r had been c a s t , and of the four young that were born, only three survived to three weeks, consequently the weights a t the l a t t e r date were high and give no i n d i c a t i o n of the weights of the "White r a t s as a group. The most outstanding e f f e c t s obtained from the colored f i l t e r s were i n the Blue, During the season 1939-40 no l i t t e r s were cast i n the Blue cage; i n the subsequent season s i m i l a r r e s u l t s were obtained. At the age of 206 days these r a t s were i n j e c t e d w i t h s y n t h e t i c .gonado trophic hormones i n an e f f o r t to stimulate reproduction. Each of tyio females received three intramuscular i n j e c t i o n s of I cc. at 3 day i n t e r v a l s , of o e s t r a d i o l benaoate i n sesame o i l (0.2 mgm./cc ); the t h i r d female was l e f t u ninfected. The male r e c e i v e d l i k e doses of testosterone propionate (5mgm/cc). These prepar-a t i o n s are known, r e s p e c t i v e l y , as Ben-ovocylin and Perandren, and were supplied w i t h the compliments of CIBA Co. L t d . , of "Montreal. These treatments, however, f a i l e d to produce r e s u l t s , so i n May 1941, a new s e r i e s of 3 r a t s were placed i n a new cage f o r observation, but at the end of 120 days, had f a i l e d to produce a l i t t e r * The greatest number of young were born under the Red l i g h t but here, where l i t t e r s were l a r g e , b i r t h weight and three week weights were low (see Table 3 ) . Percentage s u r v i v a l was s l i g h t l y below average, but the deaths, here, were more of t e n due to cannibalism than under any other c o l o r . In some instances whole l i t t e r s were devoured at a time. The next g r e a t e s t number of young were produced under the V i o l e t f i l t e r , but u n l i k e the Red, percentage s u r v i v a l was highest of a l l colored f i l t e r s , , B i r t h weight a l s o proved to be highest (equal to the b l a c k ) ; however three week weights dropped to t h i r d highest. Following the V i o l e t , the r a t s born under Green l i g h t showed the next highest b i r t h weight* and the highest three week weight. Here-* however, the number of young produced was r e l a t i v e l y s m a l l , although the percentage s u r v i v a l was. above average. Small l i t t e r s , and the subsequent b e t t e r i n d i v i d u a l n u t r i t i o n may have played a part i n p l a c i n g the three week weight so h i g h . This f a c t o r was p a r t i c u l a r l y apparent i n the case of the Bed, where l i t t e r s were, l a r g e and as a consequence, three week weights were low. Yellow'and Grange l i g h t appeared very s i m i l a r i n t h e i r e f f e c t s ; b i r t h weight i n the former was higher, but - t o t a l number born, three week weights and percentage s u r v i v a l were p r a c t i c a l l y i d e n t i c a l f o r the two c o l o r s . In general the e f f e c t s were intermediate between Green and Red as f a r as the weights were concerned. The c o n t r o l Black r a t s "Showed the highest i n d i v i d -u a l b i r t h weight and although the t o t a l number born was not very l a r g e , w i t h such a h i g h percentage s u r v i v a l (83%), the three week weight was the lowest recorded* I t was i n these p a r t i c u l a r cases that i t was found o f t e n impossible to wean the young a t three weeks; i n a b i l i t y to eat the checkers, or even d r i n k from the water b o t t l e s , r e s u l t e d i n such a weak-ened c o n d i t i o n that the mother was l e f t i n the cage f o r a week or so -23-Sex rafeio With such small groups of data on which to work, together w i t h the f a c t that in. some cases i t was found impossible to sex the young w i t h any c e r t a i n t y e a r l i e r than two weeks of age, the values recorded f o r the sex r a t i o propably bear l i t t l e significamice, as they are presented (see Table 3 ) . However i t may be pointed out that i n some cases c e r t a i n sexes appear to predominate. Thus, equal numbers of both sexes were produced under V i o l e t , •Yellow, and Red, while females predominated under Orange l i g h t and males under Black, alhaost i n the r a t i o of two to one. As many of the r a t s were not aexed u n t i l three weeks of age, during which time there was a f a l l i n g o f f i n s u r v i v a l due to m a l n u t r i t i o n or c a n n i b i l -ism, some of the values recorded do not represent the sex r a t i o a t b i r t h . Toward the l a t t e r part of the experiment the l i t t e r s were destroyed a f t e r o b t a i n i n g the b i r t h weight, i n order to* Jseep the colony down to a reasonable number; the sex r a t i o of such l i t t e r s , of course, were not determined. Reccurence of Pregnancy (See F i g . 3) I n a l l instances where t h i r d l i t t e r s were c a s t , the time e l a p s i n g between the second and t h i r d l i t t e r s was always greater than f o r that between f i r s t and second. N Greatest v a r i a t i o n f o r age a t which f i r s t l i t t e r s were cast occurred i n the V i o l e t . Here a d i f f e r e n c e of 48 days was recorded between the ages of the f i r s t and l a s t female to cast t h e i r f i r s t l i t t e r s . This value decreased i n the Green, reached a minimum of 4 days i n the Yellow and rose again to 12 days i n the Red. This same general trend h e l d good f o r the time elapsing between the f i r s t and l a s t of the second l i t t e r s cast w i t h i n the same c o l o r groups. V i o l & t again showed a d i f f e r e n c e of 77 days, Green 22 days, Yellow and Orange 6 days, and Red 12 days, whether t h i s i s an e f f e c t of the r a d i a t i o n , or whether r e g u l a r i t y i n recurrence of pregnamcy i s a. c h a r a c t e r i s t i c of an i n d i v i d u a l , i s debateable. In the case of the V i o l e t female (£) who cast her f i r s t l i t t e r a t 70 days of age, there appears to be s i g n i f i c a n c e i n the observation t h a t , although pregnancy occurred i n the normal l e n g t h of time (49 days), by the time moat of the "other females had c a s t t h e i r second l i t t e r s , she had cast her t h i r d . The same appeared i n the Red, where the $' cast her t h i r d l i t t e r at 149 days of age, but she d i d not r a i s e a l l her young to three weeks, hence she had been returned to the male sooner than u s u a l . A c t u a l shortening or lengthening of the time of recurrence of pregnancy from normal would perhaps be of more significraniee, but such observations d i d not appear i n t h i s present work. Reproduction ffi Generation (See Table 5) At the time of w r i t i n g too l i t t l e data was a v a i l a b l e from which to draw -25-any very d e f i n i t e conclusions, except t h a t , as was mentioned "before* the second group of Blue r a t s f a i l e d to produce any l i t t e r s up to 120 days of age. Siz e of l i t t e r s appeared to be f o l l o w i n g the same general trend, w i t h the exception of the Red where only 3 were born i n the f i r s t l i t t e r , none of which survived to three weeks. Age a t time of c a s t i n g of f i r s t l i t t e r s a l s o appeared to be r e c a p i t u l a t i n g the F q generation, although data f o r Orange and Red i s l a c k i n g . D i s c u s s i o n and Conclusions The r e s u l t s of previous work has "been f a i r l y w e l l v e r i f i e d as regards the l a c k of s i g n i f i c a n t d i f f e r e n c e i n the weights of r a t a r a i s e d under the various colored l i g h t s , however* d i f f e r e n c e s do seem to make t h e i r appearance when such data as "birth weight, three week weight, s u r v i v a l and occurrence of pregnancy are considered. The t o t a l absence o f l i t t e r s I n the Blue cages, p o i n t s to the f a c t t h a t the Blue cellophane e i t h e r transmits c e r t a i n r a d i a t i o n s that i n h i b i t pregnancy, or f i l t e r s out c e r t a i n r a d i a t i o n s that are necessary to stimulate pregnancy. The d e f i n i t e suppression of l i t t e r s i n the white ( c l e a r ) cellophane bears out the former p o s s i b i l i t y , since such - I n h i b i t i n g r a d i a t i o n s would tend to be transmitted by both f i l t e r s . E v i d e n t l y i t i s the blue end of the spectrum t h a t i s concerned here, as the red end i s passed i n d i f f e r e n t p o i n t s by a l l the other f i l t e r s , even i n c l u d i n g the Y i o l e t , which contained much r e d d i s h dye. There i s , however, the p o s s i b i l i t y that d i f f e r -e n t i a l r a d i a t i o n e f f e c t s may not be concerned here a t a l l , s ince i t has been shown that Blue cellophane has the property •*. P r i v a t e communication of producing a a l i g h t degreeeof p o l a r i z a t i o n of the transmitted l i g h t . This c h a r a c t e r i s t i c i s very s l i g h t , or e n t i r e l y absent i n a l l other colored cellophanes, so appears, to be more a property o f the dye incorporated i n t o the cellophane, rather than of the cellophane i t s e l f . I f p o l a r i z a t i o n i s concerned here i t i s d i f f i c u l t to understand how such a small degree, could produce such s t r i k i n g r e s u l t s . With i n c r e a s i n g wavelength a decrease i n b i r t h weight and three week weight becomes evident, u n t i l i n the Red, large l i t t e r s are produced, r e s u l t i n g i n an immature c o n d i t i o n of development a t three weeks; cannibalism seems to predominate here. Luce—Ciausen and Brown's (26) observation was not borne out i n regard to the s u r v i v a l of the young, although -they g i v e no d e f i n i t e f i g u r e s f o r the c o n t r o l s , other than r e p o r t i n g that the v i s i b l e r a d i a t i o n s promoted the s u r v i v a l . In the present work, the highest s u r v i v a l occurred i n the c o n t r o l Black, f o l l o w e d by V i o l e t . D i f f e r e n t i a l wavelengths show l i t t l e e f f e c t upon the srecurrence of pregnancy, w i t h the exception of the White, where apparently i t was i n h i b i t e d , but sexual maturity appears to be hastened to a s l i g h t degree by the longer wavelengths, e s p e c i a l l y by the Yellow-orange r e g i o n . This observation i s supported by the work of Bissonnette and others (see above), i n which spermatogenesis i n male animals i s promoted by the longer wavelengths, and which has been o f f e r e d as an explanation f o r the seasonal spermatogenesis and r e g r e s s i o n rhythms recorded i n b i r d s . Of d o u b t f u l s i g n i f i c a n c e was the observation that i n a l l cases where t h i r d l i t t e r s were cast, the time elapsing between the second And t h i r d was always shorter than that elapsing between the f i r s t and second. However, s i n c e , at the termination of the experiment, n e i t h e r Black female had cast her t h i r d l i t t e r , there i s no evidence that sucBa an e f f e e t i s due to i n c i d e n t r a d i a t i o n ; i t could q u i t e w e l l be a n a t u r a l occurrence. Although data f o r the F j generation reproduction was incomplete, sexual m a t u r i t y showed a very close c o r r e l a t i o n w i t h (fche FQ generation, and such remarks as have been made f o r the l a t t e r , hold good f o r the former. A c t i v i t y t e s t s are very i n t e r e s t i n g i n the l i g h t of the methods used f o r neuropathic cases i n asylums. Morbid, morose p a t i e n t s are placed i n orange or red rooms while the overactive ones are placed i n blue rooms. Such f a c t s are w e l l borne out i n the a c t i v i t y curves f o r the males ( F i g . 2 J , Where the Red r a t s proved the most a c t i v e of those under colored cellophanes. Blue, although not the l e a s t a c t i v e , nevertheless, shows a f a i r l y smooth curve• These Blue r a t s have gonadotrophic i n j e c t i o n s which considerably increased t h i s a c t i v i t y as observed i n t h e i r movements i n the cage. Comparison of the Black and the White points to the f a c t that l i g h t has a d e f i n i t e ^deadening'8, e f f e c t upon -29-the n a t u r a l l y nocturnal a c t i v i t y . This may w e l l he the r e s u l t of a visual-hormone r e a c t i o n , since the Blue and the White were non- and low-producers r e s p e c t i v e l y , as f a r as reproduct-i o n i s concerned. Likewise the greater a c t i v i t y of the Red may he c o r r e l a t e d w i t h h i g h reproduction. Suchobservations l e a d to the conclusion that blue l i g h t i s d e f i n i t e l y concerned i n the processes of reproductive e f f i c i e n c y , f o r both the white and the blue f i l t e r s passed the blue end of the spectrum, as has been pointed out before. H i s t o l o g i c a l examinations of reproductive organs of such r a t s would prove very i n t e r e s t i n g and should provide the grounds f o r f u r t h e r research on the p h y s i o l o g i c a l e f f e c t s of v i s i b l e r a d i a n t energy. I t i s s i n c e r e l y hoped that t h i s present work has " l a i d the foundations f o r such research, and that the d i f f e r e n t phases of the reproductive and h i s t o l o g i c a l r e a c t i o n s w i l l be studied i n more d e t a i l a t some f u t u r e date. ITIstar a l b i n o r a t s were bred and r a i s e d under f i l t e r s of v i o l e t , Blue. Green, Yellow, Qrange, Bed, Clear and Black cellophane f o r one season of four months and f o r another of eight months. S i z e and weight of l i t t e r s , together w i t h other data p e r t a i n i n g to reproduction, as w e l l as weekly weight and a c t i v i t y have been recorded. Blue l i g h t i n h i b i t s sexual maturity to the; extent that no l i t t e r s were cast d u r i n g the e n t i r e period of the experiment. White l i g h t suppresses reproduction, while the l o n g e r ato&$m«£ wavelengths appear to hasten sexual m a t u r i t y , although they are accompanied by a lower b i r t h and three week weight "due to l a r g e r l i t t e r s being produced. Recurrence of pregnancy and general body weight are unaffected. A c t i v i t y i s markedly higher i n Black and Red, than i n White or other colored r a d i a t i o n s , and i s greater f o r males than females. Bibliography I . Browman, L 4 G . , Jour. Exper. Z o o l . 74, 375. 1937. 2v Brown, W.H., Jour. Exper. Med. 48, 31. 1928. 3. Brown, W.H., Jour. Exper. Med. 48, 567. 1928. 4. Brown, W.B., Van A l l e n , CM., Jour. Exper. Med. 42, 69. 1925. 5. Bissonnette, T.H., P h y s i o l . Z ool. 5, 92. 1932. 6. Bissonnette, T.H., Wilson B u l l . 51, 227-232. 1939. 7. Coblentz., I.W., Carnegie I n s t . Pub. 164. 1912. 8. C o l l i e r , H.B. and Wasteneys, H i , A u s t r a l i a n Jour. Exper. B i o l . Med. S c i . 9» 89-112. 1932. 9. C r o z i e r , W.J., Jour. Gen. P h y s i o l . 23, 3G9-320. 1940. 10. Del C a s t i l l o , E.B. and P i n t o , A., Rec. Soc. Argentina B i o l . 5, 463-469. 1939. 11. Dessauer, F., Radiology 24, I-I4. 1930. 12. B e t w i l e r , Jour. F o r e s t r y , 29, I3I-I33. 1931. IS. Duggar, B.M», The B i o l o g i c a l E f f e c t s of R a d i a t i o n . 1.4. Emmons, CW. and Hollaender, A., Am. Jour. Bot. 26, 467. 1939. 15. F a i r h a l l , L.T., Am, Jour. P h y s i o l . 84, 378. 1928. 16. P u l l e r s H.J., Am. Jour. Bot. 18, 895. 1931. 17. Gates,'E.L., Jour. Gen. P h y s i o l . 13, 249. 1929. 14, 31. 1930. 18. H a r r i s , G.W., Proc. Boy. .Soc. B. 112, 374. 1939. 19. Hamley-Gartwright, Q., Z e i t . Phys* 9Q, 482. 1934. 20. Heyroth, The Chemical A c t i o n of U l t r a - v i o l e t Rays. 21. H i l l , M. and Partes, A.S., Proc. Roy. Soc. B. 113, 537, 1933. 22. H i l l , M. and Parkes, A.S., Proc. Roy. Soc. B. 115, 14. 1934. -23. Landen, E.W., Am. Jour. Bot. 18, 145. 1938. 24. L a v i n , G.I. and Stanley, W.M., Jour. B i o l . Chem. 118, 269-274. 1937. 25. Luce-Clausen, E.M* and Brown, E.E., Jour. N u t r i t i o n 18, 537-549. 1939. 26. Luce-Clausen, E.M. and Brown, E.E., Jour. N u t r i t i o n 18, 551-562:. 1939* 27. M a r s h a l l , E.H.A., Jour. Exper. B i o l . 17, 139. 1940. -32-28. M a r s h a l l . G.H.A. and Bowden, P.P., Jour. Exper. B i o l . I I , 409. I934o 29. Matthews, L..H., Proc, Roy. Soc. B. 126, 557. 1939. 30. McAulay, A.L.. and Taylor, M.C., Jour. Exp. B i o l . 16, 474. 1939. 31. Meier, P., Smithsn, Ann. Rep. 359. 1933. 32,. Hurnberger, G.E., Proc. Nat. Acad. S c i . 23, 189. 1937. 33. Packard, Ann;. Rev. P h y s i o l . 101, 29. 1939. 34. Pearce, L., and Van A l l e n , CM., Jonr. Exper. Med. 44, 447. 1926. 35-. P r i c e , W.G. and Gowan, J.W., Phytopath. 27 , 267-282. 1937. 36. Ramsey, G.B, and B a i l e y , A.A., Bot. Gaz. 89, I I S . 1930. 37. Sheard, G. and Higglns* G*H«., Am. Jour. P h y s i o l . 85, 290* 1928. 38. Sheard, G. and Higgins, G.M., Am. Jour, P h y s i o l , 85, 299. 1928, 39. Wlthrow, R.B. and Benedict, H.M.,, B u l l . B a s i c . S c i . C i n c i n n a t i 3, I6 I - I 7 5 . 1931. 40. Zahl, P.A., E o l l e r s L.R. and Haskins, C P . , Jour. Gen. P h y s i o l . 22, 689. 1939. TABLE •ir-s WEIGHT AKD UTTER RECORDS. FQ GENERATION. AGE 'WHITE i 6 o cf' „ BLACK 6 Q JIOLET d Q Q' Q" Cf BLUE _Q _ Q' Q" GREEN ; Cf Q 9' YELLOW -: Cf Q Q' ' ORANGE : Cf 0 Q' RED ; CT Q Q1 0" 21 33 32 34 25 24 29 21 26 20 27 22 21 20 19 19 26 26 20 19 18 34 19 18 ; i 1 : 3 ! 34 34 29 30 28 52 49 54 24 38 38 40 37 36 27 : 19 27 23 26 24 54 22 24 52 50 35 57 50 54 49 59 61 63 61 65 60 47 37 49 55 45 48 43 41 42 56 56 42 108 95 114 82 73 75 72 70 81 76 88 78 78 71 80 114 113 96 77 80 ' 49 135 109 138 106 93 89 99 85 100 96 120 98 103 94 : 90 90 104 ; 92 87 83 141 84 82 139 111 96 101 56 **58 118 160 136 114 109 122 109 117 120 148 114 116 113 111 109 118 111 101 99 165 100 98 165 126 116 117 181 128 180 145 112 116 129" -145 137 126 145 136 113 119 178 115 112 190 138 132 135 70 202 136619.6; 172 130 139 187 139 155 145 Litter ^ 70—11—54. gr. 201 146 137 144 158 140 155 161 128 129: 199 124 119 214 153 143 145 77- 227 153 213 189 144 146 308 152 171 157 218 159 145 151 180 146 159 181 138 152: 216 133 •130'! 230 156 160 153 84 230 153 228 208 151 160 223 160 180 172 233 165 151 160 191 164 174 194 1 59 160: 232 154 156: 238 163 155 160 87 149 150 168 155 160 91 245 172 238 208 160 156: 237 142 189 183 246 172 158 166 205 173 185 202 164 179: 240 174 163! 255 179 167 172 93 173 167 162: Litter. 2" 92—3—17 gr. 200 180 179 172 3 98 259 184 246: 226 180 175: 242 151 187 188 258 186 162 174 208 190 213 Litter. 100-6-32 gr 208 198 197s Litter.? 100-5-27 gi 247 205 179s 266 197 183 173 101 199 192 184 180 159 210 217 185 227 195 183 Litter TABLE 1 HEIGHT AND LITTER RECORDS. FQ GENERATION (Cont'd.) AGE i „ WHITE € Q cP' Litt e r 9. 104-4^18" gr BLACK 6 Q Q' VIOLET 6 Q Q' .0" :• n BLUE c? Q p' q : GREEN <S Q Q' YELLOW Cf Q Q1 Litter 2' 104-7-33 gr ORANGE C? Q Q' Litter ? 102-9-37 g ^ RED C 0. 0' 0" • 105 249 176 242 231 212 189 Litter $ 108-9-48 gr 243 171 205 194 :272 183 183 196 :210 205 201 207 181 180 250 170 208 Litter *' 108-7-35 g 270 198 207 181 Litter 9-IO5-9-46 gr. Utter $< 1 0 5 ^ 3 7 gr ,108 2©' 188 195 112 273 202 255 234 182 222 L i t t e r $' II4-7-35 gr 262 195 224 210 283 201 177 183 219 198 220 Litter $ 115-3-17 gr 223 202 197 262 177 184 274 216 156 209 • 115 Litter $' 118-6-37 gr 222 Litter 117-10-43 gr 119 218 264 241 194 197; 265 186 213 19$ Litter £ 119-10-^6 gr. 293 215 184 195 234 208 208: 237 202 172; 268 178 199 283 212 183 191 122 224 269 192 215 186 194: 192 126 251 176 201: 263 188 216 185 300 207 192 193: 242 215 197: 227 196 204 271 189 200 272 200 184 183 129 207 206 208 210 178 194 184 210 198 198 WEIGHT AND LITTER RECORDS. F Q GENERATION (Cont'd.) AGE i WHITE 6 Q 6' BLACK Cf Q 9' !' VIOLET O Q CO1 Q" BLUE S Q' GREEN Cf Q 9' . YELLOW : ORANGE d 9 Q' : . RED ( T O 0' 0" 133 194 272 >69 203 216 201 i-302 219 189 200 235 191 247 215 193 :28l I87 190 [278 205 136 195 187 207 ! 184 217 218 2ol 201 204 186 225 200 ! 190 183 : 192 211 191 140 249 169 179 i290 172 214 216: 310 220 193 200 240207 186 252 234 207 \ 208 194 : 197 192 143 200 187 194 202 245 Litter 9" 144-4-20.5 gr 228 197 209 198 209 266 218 Litter ? 146-14-79 g] \ 216 199 : 211 243 133 DIED Litter £' 146-11-53 gr 147. 183 270 117 189; 287 189 211 217 310 233 202 201: 2l6i 288 :28l 199 204 150 209 191 193 170 220 192 228: 245 253: Litter ?' l52-9-48~gr 222 245 Litter 9' 152-9-39 g: : 229 191 • 154 ' 200 289; 265 201 199 '272 159 229 216 : 312 224 197 215s 223 188 Litter 188: ' gr 254 218 210; 243 210 ! 2.13 157 210 : 266 212 204: 170 237 227 i 228 206 217; 212 209: 233 200: 271 215 280 22.7 iee 158 237 233 200 Litter 158-10-54 t IT 199 287i 270 242 215? 264 177 228 230 s 328 227 215 216: 246 214 223: 250 212 210: 212 171 287 176 163 200 227 s 227 Litter 9-165 199 204 243 181 225 227 207 220 216 231 207 TABLE T WEIGHT AND LITTER RECORDS. F 0 GENERATION (Cont'd.) 205 286 211 302 21? 216 315 205 215 316 211 215 316 VIOLET cf $ ip 2" |: 267 216. 210L 188 209 Litter ?' Litter « 168-9-43 gp l£S^p?3 gr 267 221 221 264 188 227I282 236 222 207 175 215 267 158 198 262 169 187 258 175 176 173 184 270 196 197 BLUE 0 g $' 2" 244 212 208 279 217 233 208 198 241 208 258 190 219 211 265 181 244 212 264 172 212 211 Litter ?' 196-12-57 gr 234 218 225 330 237 217 232 235 215 231 312 234 218 226 244 222 233 314 242 214 230 241 216 223 326 246 211 228 GREEN 0 $ oj 236 214 197 227 188 242 248 179 Litter £ 176 5^-24 gr 228 192 241 235 194 241 200 242 250 201 201 227 248 205 :257 179 202 b YELLOW cf 2 2' k245 193 200 ORANGE <r 2 2' 243 213 193 183 204 255 196 201 206 206 260 209 208 208 231 262 212 228 . RED .& 2 9' 233 184 225 200 188 215 158 201 176 206 185 204 196 206 204 214 207 222 299 204 178 194 183 290 183 191 294 180 202 292 176 215 182 218 293 186 232 196 258 Litter £' 195-10-49.5 gr TABLE 1 ' WEIGHT AND LITTER RECORDS. FQ GENERATION (Cont'd) WHITE 6 9 d" 213 218 316 214 202 199 270 210 209 227 317 BLACK 232 217 :271 201 230 166 246 214 265 170 230 201 218 182 216 194 220 275 189 205 213 241 210 235 324 248 216 227 250 218 232 321 251 204 225 „ GREEN 2o6 269 253 211 212 215 209 257 218 208 YELLOW :<r q 9 216 246 260 215 26? Litter 9' , 2 0 5 ^ 3 9 . 5 gr 217 228 264 227 235 204 , Litter * , 212^29.5 gr ORANGE tf 0 9' 199 231 208 273 224 Litter 20W^52 gr. , RED (T 9 9' 300 212 222 222 212 296 229 196 242 194 213 221 288 267 179 ,1 TABLE 2 INDIVIDUAL LITTER RECORDS. F 0 LITTERS. LITTER I NUMBER BIRTH WEIGHTS VIOLET 9- 1 2 3 9 ' 1 2 3 4 2 In toto 11 10 9 6 5 12 6 3 4 54.0 56.0 53.0 37.0 57.0 35.0 17.0 20.5 I n d i v . 4.92 5.4 5.89 6.17 4.96 5.84 5.67 5.1 NO. SURVIVING TO 3 WEEKS 8 10 9 6 10 3 4 3 WEEK.WEIGHTS In toto 91.0 186.0 193.5 117.0 173.0 105.0 152.0 Individ. 11.4 18.6 21.5 19.5 17.3 35*0 38.0 STATISTICAL MEANS Indiv.birth wt. ra= 5.45 Indiv.3 wk.wt. m= 23.16 GREEN 9 1 2 3 o_-i-YELLOW 9 1 2 3 3 5 7 6 6 5 14 5 17.0 24.0 44.2 32.0 27.0 5.67 4.8 6.27 5.34 4.5 2 5 4 6 66.0 149.5 130,0 181.0 33.0 29.9 32.5 30.2 m= 5.25 27.0 79.0 29.5 5.4 5.6 5.8 2 12 82.0 145.0 41.0 12.1 TABLE 2' INDIVIDUAL LITTER RECORDS. F Q LITTERS. (Cont'd) LITTER NUMBER BIRTH WEIGHTS NO. SURVIVING 3 WEEK WEIGHTS • STATISTICAL MEANS m to to Indiv. • TO 3 WEEKS In toto Individ. Indiv.birth wt. Indiv. 3 wk. wt. YELLOW $' 1 7 33.0 4.72 4 108.0 27.0 2 9 48.0 5.34 5 146.0 29.2 3 8 39.5 4.87 8 175.0 21.9 m = 5° 11 m = 27.25 ORANGE 9- 1 9 37.0 4.11 6 118.0 19.6 2 10 54.0 5.4 7 154.0 22.0 7 35.0 5.0 5 123.0 24.6 2 9 39.0 4.34 ^ 2 44.0 22.0 3 9 52.0 5.78 m=5.o5 m= 21.9 RED $ 1 9 46.0 5.11 8 146.0 18.25 2 11 58.0 5.28 11 181.0 16.5 3 12 67.5 5.64 0_« 1 8 37.0 4.63 0 2 11 53.0 4.83 7 •1.61.0 23.0 3. 0J< 1 10 10 49.5 43.0 4.95 4.3 10 4 179.5 95.5 17.95 - 23.9 m= 4.87 ra= 19.95 INDIVIDUAL LITTER RECORDS. F Q LITTERS. (Cont'd) LITTER NUMBER BIRTH WI SIGHTS NO. SURVIVING 3 WEEK .WEIGHTS'." •STATISTICAL MEANS In toto Indiv. TO 3 WEEKS In toto Individ. Individ.birth wt. Indiv. 3 wk. wt. BLACK 9- 1 9 48.0 5-34 ; ' . 9 I63.O 18.1 2 10 52.0 5.2 10 202.0 20.2 6 36.5 6.09 ' ; J ' 1 7 35.0 5.0 6 " . I8O.0 30.0 2 43.0 4.78 9 193.0 21.5 ••' WHITE 9 1 4 18.0 '4.5 3 99.0 : 33.0 roi t-H EHl CO M CO 1-3 <d B c« EH EH M t-H 1 o fx) ro <t! o EH EH B co ro ojb © M . EH CO «aj B -, w 1-3 o «i PQ EH o • EH o B O 1-3 o o XT\ CM H ro CM H Cvl Cvl EH l-H o > o ON vO ft • • ro OX O XTN ro XTs ro CO CM ro Irs CM CM o CM Xrs H CO O • CO XTS Xrs H Xf\ ITS cvl H O 00 ITS • • • ft • • Xrv Xrs ITS Irs Irs Xrs Xrs ON CM ON • « • • • H H H H O H Xrs Xrs 00 - H CM H 00 CM co o ro CM Os CO H ON CM CO CM t-t -4-H ro ro CM CvJ O ro B PC o hi hi B O Xrs O CM ro B o B *3' '* i Q w Q o EH •3 H PQ m PQ T A B L E 4 • WEIGHT AND L I T T E R R E C O R D S . F X GENERATION AGE s WHITE (?' J B L A C K C? Q 9' 1 V I O L E T C? Q 9. ! B L U E cf 2 9' ! . GREEN cf 2 9. ! JffiLLOW ct 9 9' : ORANGE Cf - Q 9' cf 0 cf' 21 s 33 33 33 i 18 18 18 : 26 28 26 ! 32 31 32 : 42 40 20 : 20 20 20 . 18 18 18 28 57 50 60 • • - 1 53 56 50 ; 49 46 48 73 65 49 29 26 26 21 21 30 : 40 37 34 35 s 56 54 48 » 78 82 69 I 76 72 96 85 71 : 48 51 50 38 i l06 88 101 • • 90 51 42 112 101 119 1 81 76 68 1109 106 90 : 108 69 67 48 45 :101 105 85 * 1 ! 117 117 107 88 88 49 115 116 127 i l28 116 105 :139 131 122 152 118 i l l :100 105 99 88 92 80 52 109 97 t • 56 134 116 143 il24 120 104 il63 138 1241 173 126 122 ;121 116 109 121 109 107 63 166 135 176 • :183 145 133 5179 145 142s 183 128 137 ,150 133 137 154 133 138 70 183 164 204 5 205 160 147 t212 173 142J 210 147 149 172 143 149 173 152 159 73 170 5154 141 123 151*. 164 74 168 159 157 148 150 76 149 131 220 178 77 193 180 215 219 178 145 152s 226 168 165 189 152 159 192 170 177 80 175 155 133 179 174 170 156 156 8l 198 • • 152 166 TABLE 4 WEIGHT AND LITTER RECORDS. F± GENERATION (Cont'd) JHHITE <5 9 c f 202 207 219 Litter $ 8 5 ^ 1 8 7 5 g 183 197 195 217 199 210 203 233 204 217 217 242 BLACK <? 9 Q-t VIOLET cf 9 9' 165 130 195 172 138 174 147 209 179 154 181 156 213 186 168 207 200 206 166 L i t t e r 2' 106-6-34 gr 193 154 228 207 159 234 160 225 259 162 Litt e r £ 97-13^9 glp 206 163 232 192 164 177 167 * BLUE <$ 9 9' GREEN j CT 9 g' 232 179 160 178 165 232 181 176 183 183 238 192 198 192 216 L i t t e r 5' 103^5-25.5 245 204 185 YELLOW ; Cf 9 9' 233 172 167 178 172 161 163 187 181 211 165 164 193 185 165 168 224 177 169 213 196 ORANGE cf 9 9-203 159 163 214 168 172 168 170 246 175 216 Litter 9 105- 10-53 gr 191: Litter 2' 106- 4-24 gr 220 169 169 RED & 9 Cf' 200 169 183 182 215 189 204 195 225 202 212 225 230 229 220 206 t i t t e r ? 106-3-19 gr TABLE 4 WEIGHT AND LITTER RECORDS. F± GENERATION (Cont'd) AGE ! WHITE 6 g c f ' s BLACK Cf Q Qt : VIOLET (5* Q Q1 : „ BLUE : „ GREEN o* 5 Q : YELLOW" c? 9 Q' ; ORANGE d1 9 Q' • RED (f 9 c f 1 109 212 :217 215 160 :233 164 168 210 186 : 197 196 172 173 191 110 213 112 228 202 251 196 171 Litter ? 112-8-45.5 ! ;r 242 219 241 214 209 225 175 176 235 200 229 115 20? 222 206 184 210 116 252 154 172 242 Litter £ 117-5-28.5 J 213 205 IT 174 174 119 237 214 262 212 184 158 175 238 184 205 229 173 176 ;243 209 236 O CO H (xj > H > Bro| CO d o £5 EH CM rH. UN NO CM CM CO CO E-i O M C£3 Individ. ro • H 28.0 O * ON ro CM ro CM • UN CM • H ro 33.7 CO • ro CM Eg 3REE WEE] o -p o p d l ,,t \D ON rH o H O 9 ON ro O • H ro H UN • • H UN H < UN • ro \o UN 0 NO O # rH ON H (—*i EIGHTS: > •H •a. i, i ro • UN UN CM • . • UN O • • SO ro • UN ro vo H • NO • UN • UN p-1 « EH o p o p £} M O • ON NO UN • UN CM UN e CO CM O a CM O « ro UN o • ON H UN • CO H o " • ro UN • UN NDlffiER ro •H NO UN O rH ro NO CO EH... EH M (-3 EH i-^ o+ o o -l-H of o+ O o+ EH o -M Of «aj 0+ 0+ l-H pq T H E H U O H f l O W E N S C O . , L I M I T E D N o . S I S B l O L I N E S 1 I N C H iaz-11-a* 

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