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Chemical study of the incipient decay of red alder, Alnus rubra, Bong. Cserjesi, Antal Janos 1961

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• CHEMICAL STUDY OF THE INCIPIENT DECAY OF RED ALDER, ALNUS RUBRA, BONG. by ANTAL JANOS CSERJESI Forest engineer, University of Forest Engineering, Sopron, Hungary, 1950 A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF FORESTRY in the Faculty of Forestry We accept this thesis as conforming to the required standard THE UNIVERSITY OF BRITISH COLUMBIA March, 1961 In presenting t h i s t h e s i s i n p a r t i a l f u l f i l m e n t of the requirements f o r an advanced degree at the U n i v e r s i t y of B r i t i s h Columbia, I agree tha t the L i b r a r y s h a l l make i t f r e e l y a v a i l a b l e f o r reference and study. I f u r t h e r agree that permission f o r extensive copying of t h i s t h e s i s f o r s c h o l a r l y purposes may be granted by the Head of my Department or by h i s representatives. It i s understood that copying or p u b l i c a t i o n of t h i s t h e s i s f o r f i n a n c i a l gain s h a l l not be allowed without my written permission. Department of The U n i v e r s i t y of B r i t i s h Columbia, Vancouver 8, Canada. Date i i ABSTRACT The chemical properties of the wood of red alder (Alnus rubra Bong.) were examined after exposure to decay by three organisms, Fomes  pinicola (Schwartz ex Fries) Cooke, as a brown rotting fungus, Polyporus  versicolor L. ex Fries and Stereum hirsutum Fr. as white rotting fungi. Cold- and hot-water solubility were most affected by all three test fungi. The increase in methanol-benzene solubility and in one-per-cent sodium hydroxide solubility were the highest in samples decayed by F. pinicola, but the changes were less pronounced than observed for the water solubles. Destruction of cellulose and lignin appeared in samples decayed by P. versicolor. The deterioration of cellulose and lignin by the other two fungi was within the limit of experimental error of the determination methods used. CONTENTS Page Abstract Introduction 1 Literature Review h Materials and Procedure 13 Results . . . . . . . . . 18 Discussion . . . . . . . . . . . . . . . . . 20 Conclusions . 23 Summary • 21+ Literature Cited 2$ Appendix I (Tables) 31 Appendix II (Figures) . . . . . . . 36 i v LIST OF TABLES Table Page I, Percentage of cold-water, hot-water, methanol-benzene and a l k a l i solubles, holo- and alpha-cellulose and lignin i n sound and decayed wood samples. (Per-centage expressed on the basis of the weight of decayed wood) 32 I I . Percentage of cold-water, hot-water, methanol-benzene and a l k a l i solubles, holo- and alpha-cellulose i n alder wood. (Percentage expressed on ttie basis of the weight of original sound wood) 33 I I I . Values of weight losses and incubation periods i n the preliminary experiment with P. versicolor . . . . . 3 4 IV. Values of weight losses and incubation periods i n the preliminary experiment with S. hirsutum 35 V LIST OF FIGURES Figure To Follow Page 1. Average annual depletion i n immature and mature forests of British Columbia . . . . . 3 7 2. Comparison of annual decay loss with annual growth i n the immature and mature forests of British Columbia 37 3 . Butter dish as incubation chamber . . . . . . . . . . . 3 8 U. The calculated curve between incubation period and weight loss caused by P. versicolor 39 5. The calculated curve between incubation period and weight loss caused by S. hirsutum liO ACKNOWHEDGMENT Sincere thanks are extended to Dr. R. W. Wellwood, and Mr. R. ¥. Kennedy of the Faculty of Forestry, and Dr. J . E. Bier of the Department of Biology and Botany, University of B r i t i sh Columbia, for their guidance, assistance and encouragement throughout the study. The help of a l l others who assisted i n the various phases of this project i s gratefully acknowledged. INTRODUCTION The degradation o f wood by microorganisms i s of great importance. I t i s b e n e f i c i a l as a source of humus i n the s o i l , but on the other hand causes a great l o s s as a causal agent i n the destruction of wood products. From the b i o l o g i c a l viewpoint, the d e t e r i o r a t i o n by microorganisms of a l l dead organic materials to simple components const i t u t e s one of the necessary l i n k s i n the carbon and nitrogen c y c l e , which i s v i t a l to a l l l i f e . The decay of wood can be accepted as one stage of t h i s process (UU). Decay f u n g i , which belong mostly t o the Basidiomycetes, cause the d e t e r i o r a t i o n of wood and wood products, and m i l l i o n s of d o l l a r s are l o s t each year because of t h i s d e s t r u c t i o n . A survey o f wood depletion i n the l a s t 350 years i n the United States shows forty-one per cent of the depletion of wood was caused by disease or f i r e , while twenty-one p e r cent was used f o r wood products, f i f t e e n per cent for f u e l , and fourteen per cent was destroyed to c l e a r land ( l ) . In B r i t i s h Columbia fungi are responsible f o r $1.2 per cent of the depletion (1.081 b i l l i o n cubic feet) i n the I n t e r i o r , and 11.1 per cent of the depletion ( l . l i f l b i l l i o n cubic feet) i n the Coast Region. This l o s s occurs mainly i n the most valuable, large, over-mature t r e e s ( F i g s . 1 and 2) ( 9 ) . In B r i t i s h Columbia the main f o r e s t tree species are co n i f e r s (9) , but i n Canada as a whole, twenty-eight per cent of the fo r e s t i s hard-woods (53). In other p a r t s of the world the r a t i o i s even higher f o r hardwood sj f o r example i n the United States i t i s forty-eight per cent, and i n Western Europe f i f t y - f o u r p e r cent (53) • 2 According to wheeler (67), the u t i l i z a t i o n of conifers i:s higher i n ratio to hardwoods than the ratio between them i n the forest, which selection may operate against the natural balance i n the forest. Recently hardwood consumption has increased at a rate faster than that of softwood consumption (23). Red alder (Alnus rubra, Bong.) i s one of the most important hard-wood species on the Pacific Coast, and i t i s the only species of alder i n Canada of significance commercially (llj.,66). Cottonwood, aspen and birch exceed the volume of alder i n B r i t i s h Columbia forests (9.66), but in Washington State seventy per cent of the hardwood i s alder (20). The outstanding use of red alder i s for furniture. Besides being used for furniture, red alder i s used in small quantities for fixtures, general millwork, and handles. Lately red alder has become an important pulpwood source i n the Pacific Northwest (U,19>55«6*>). The wood has also proved suitable for the manufacture of the wooden parts of shoes (21). In the forest the practice of interplanting alder in conifer plantations to improve growth of the conifers has long been recognized by Europeans and i s also practiced extensively i n Japan. I t has proved bene-f i c i a l i n Alaska also, where an average of f i f t y - f i v e pounds of nitrogen accumulation per acre under alder thickets has created a favorable site for Sitka spruce, which succeeded the alder (1*3). I t s rapid growth even on poor sites (provided they are moist), i t s soil-enriching a b i l i t y , and the increased use of the wood i n recent years, indicates that consideration should be given i n forest management plans to i t s continued production on certain types of forest land (20). 3 Red alder matures in about f i f t y years and i s considered old at eighty years. Mature trees are from 80 to 100 feet high and from fourteen to eighteen inches in diameter. Mature alder trees are subject to attack by fungi, but these cause l i t t l e damage to trees less than f i f t y years of age (65). The present study i s concerned with the incipient decay of red alder. This wood species was selected because of i t s growing importance, and because i t s decay has not yet been studied intensively. Test fungi were selected from the common species responsible for the destruction of red alder wood i n Br i t i s h Columbia. LITERATURE REVIEW The • u t i l i z a t i o n o f wood attacked by fungi causes problems i n the wood-using i n d u s t r i e s . In sawn lumber the attack o f mould, sta i n and decay fungi can be serious. A l l of them a l t e r the p h y s i c a l , physicochemical, and chemical p r o p e r t i e s of wood. Even the surface growth o f moulds causes change i n the absorption o f water i n wood. (k8,59,64). Discoloration caused by any f u n g i , even i f not accompanied by measurable strength l o s s , i s un-des i r a b l e ( 7 , 8 , 5 7 , 5 9 ) . The appearance of the wood may be made unsightly so that lumber intended f o r high-quality f i n i s h stock i s no longer s u i t a b l e f o r t h i s use. In the pulp and paper industry, decayed wood i s more widely used than i n saiamills, but the extent of decay governs the economical u t i l i z a t i o n of wood ( 5 6 ) . The development of decay can be d i v i d e d i n t o d i f f e r e n t stages. In the i n c i p i e n t or e a r l y stage the wood i s as f i r m and hard as the sound wood. The colour of the wood changes more or l e s s i n some cases, but i n other cases t h i s stage can be detected only by microscopic examination f o r the presence of fungal mycelium i n the wood. Incipient decay can be danger-ous i n c e r t a i n types o f wood u t i l i z a t i o n ; f o r example, there i s a s i g n i f i -cant l o s s i n strength i n the i n c i p i e n t areas of decay i n the case of destructive rots ( 6 2 ) . Cartwright, Campbell and Armstrong (16) found only 1,8 per cent decrease i n modulus of rupture i n ash wood decayed by Poly- prous hispidus, but f i f t y per cent drop when the wood, was attacked by Trametes s e r i a l i s . In both cases the weight l o s s was 1 .4 per cent. In the progress of decaying, the wood changes from apparently sound wood to completely decayed wood, while passing through the following stages: i n -cipient decay, intermediate decay and, f i n a l l y , advanced decay. A l l pulpwood, owing to the fact that fungus spores occur prac-t i c a l l y everywhere, i s usually more or less already infected when i t arrives at the wood-yard (5>,5'Ii)» Efforts should therefore be mainly directed to counteracting the favourable conditions for the development of decay fungi (U6,*>7), The most important conditional factors influencing fungal attack and development in wood are the moisture content, temperature and availa-b i l i t y of a i r . For the growth of fungi at least twenty per cent moisture must be present i n wood, but i f the logs are stored in water, the lack of oxygen i s the most important inhibiting factor for the wood-rotting fungi. The temperature requirements are different for different fungi. Several of them are s t i l l able to grow slowly at a temperature close to the freez-ing point. For others the temperature optimum i s above 30°C, The most common optimal temperature i s between 20 and 30°C, (62), According to Bjorkman (5) decay in pulpwood appears to be a pro-blem during storage only after the second year, but others found even oak heartwood started to rot within six months (28,31,li5>). In an experiment in the United States (62) the period marking the appearance of decay varied from six months to two years, according to the storage conditions. While lack of water or a i r stop completely the growth of fungi, cold weather just slows down the decay, owing to the inhibition of microbiological growth at low temperature. Six months storage i n the summer months usually results i n a loss in kraft pulp y i e l d of about three per cent on an oven-dry wood basis. Pulp strength, as well as yield, i s noticeably reduced by two months 6 storage i n the summer and five to six months'storage i n the winter in the southern United States (15). Therefore most southern mills t r y to l i m i t storage to th i r t y to sixty days i n -the summer months. By comparison, i n the northern states and eastern Canada, pulp m i l l s usually keep at least a year's supply of pulp wood i n storage (2k,35), where during the winter losses are practically n i l . But according to other data (15), losses i n wood i n northern United States may run up to eighteen per cent for a fifteen-months storage period, or a reduction of screened-pulp yield from about k6 to about 39 per cent. Another factor affecting deterioration of wood appeared i n a pre-liminary study carried out by Wright ( 7 0 ) . He sampled and examined outdoor chip p i l e s containing Douglas f i r and alder wood chips in different ratios. I t was found that mixing of different species (soft- and hardwoods) helped to inhibit deterioration to a certain extent i n outdoor chip p i l e s . Such large amounts of rotten wood are available for the pulp i n -dustries that i t made necessary the study of the usefulness of decayed wood for pulp making (56). Decayed wood i s undesirable for pulping since i t has a lower density, results in lower yields of pulp, and produces pulp having lower strength, darker colour, and lower alpha-cellulose content than that from sound wood. The sulphite pulping i s seriously affected by decayed wood, but this depends upon the type of decay and i t s extent. Decay by wood-destroying fungi i s serious i f i t i s far advanced, resulting i n a weak, d i r t y pulp with low brightness, and a loss in y i e l d . In the sulphate process yields are reduced, more al k a l i i s consumed, and pulp strength (bursting and tearing strength) i s lowered when decayed wood i s used. In 7 the mechanical process, decayed wood results i n lower yields, poorer colour, reduced strength, increased fines, and poorer running properties on the paper machine (!?,l5,3li,50,62,68). Decay does not affect the grinding pro-perties unless i t has reached excessive levels, i n which case i t may reduce the power required i n grinding. Most groundwood mills carefully c u l l out any decayed wood (16,62). The loss i n pulp y i e l d and pulp quality resulting from decayed wood i s thus related to the extent of decay. Weight loss i s the most commonly used expression for the extent of decay, but for i t s calculation the density or weight of the original sound wood i s necessary. In practice, however, only the already decayed wood i s available, and because of the great variation i n specific gravity even i n one tree, the stage of decay cannot be expressed by the weight loss. In this case the one-per-cent a l k a l i solubility test (69), or the combination of a l k a l i s o l u b i l i t y and alpha-cellulose content (l">), seems to be the most useful indication of the extent of decay caused by brown-rot fungi. Brown-rot i s the most common in softwoods (15.17). Kawase (I4.O) found that a l k a l i consumption by wood can be used for determination of the stage of decay. Very early stages of decay do not show considerable weight lossj or, as several workers found, the microorganisms may produce weight gain (25>,3U,UO). Therefore, for an expression of the extent of fungal attack at the early stage, strength properties of the wood have been used as indicators (36,Ul,f>8). I t i s commonly known that decayed wood i s softer and less strong than sound, and that wood in an advanced stage of decay crumbles away at touch. In an early stage of decay the effect of fungal 8 attack varies greatly according to the species of fungus involved. Though the disintegration and softening brought about by a white rot i s usually less pronounced in the earlier stages than that brought about by a brown rot, the effect on the toughness or shock resistance may be immediate, and be used to indicate incipient decay (U7,50,60). Gartwright and Findlay (17) made the following and most probable conclusions about the effect of fungi on the strength properties of w>od: (l) fungi causing brown rots bring about a fairly rapid drop in strength pro-perties of woodj (2) fungi causing white rots may also, in the case of certain species, bring about a rapid drop in toughness, but probably less rapid than by brown rotsj (3) toughness i s most rapidly affected by fungal infection, followed in approximate order of susceptibility by bending strength, compressive strength, hardness and elasticity. The chemical change in •wood components as fungal attack advances i s interesting for two reasons. Firstly, the primary effects of micro-organisms appear in the different components of wood and through these changes the mechanical properties of wood are altered. Secondly, pulp mills are very much concerned about the chemical composition of their raw mater-i a l s . Early chemical analyses of decayed wood were made mostly to examine differences between the components of sound wood and decayed wood at an advanced stage (k9). No special attention was paid to the change during the progress of decay and in several cases the species of fungi were not determined (6,27,37). In these experiments the stage of decay i s expressed differently by different workers. The most common expression i s 9 weight loss, which i s used regardless of time necessary to reach i t (2,10, 11,58). Lyr and Ziegler (U9) used the period of incubation, while Kawase (kO), i n several cases i n his study, just described the stage of decay in terms of gross appearance. The presence of fungal mycelium makes d i f f i c u l t the analysis of decayed wood. While some fungi produce relatively l i t t l e mycelium i n proportion to the weight of wood they consume, others produce considerable amounts. In a study of Campbell (12) one-fifth of the weight loss (10.6 per cent) of the original wood could be accounted for as the mycelium of Ganoderma applanatum growing on the exterior of beech wood. The mycelium of wood-rotting fungi may contain, according to the species, from three to f i f t y - f o u r per cent of a lignin-like substance. I t has been shown that sixty-one per cent of the dry weight of the mycelium of Phellinus cryptarum i s resistant to the action of the chemicals used for cellulose determina-tion and i t i s therefore considered as cellulose (13). Determination of the amount of mycelium present i n decayed wood i s f a i r l y d i f f i c u l t . I t might be estimated by nitrogen determination, because a considerable portion of the wall of most fungi consists of c h i t i n , which contains 6.9 per cent nitrogen. In most of the experiments the presence of the fungal mycelium in wood i s neglected i n the chemical analysis (22,51,69). The difference i n a b i l i t y to deteriorate wood components i s the basis of separating brown-rotting and white-rotting organisms from each other. Brown-rotting fungi destroy mostly the carbohydrate components of wood, while white-rotting organisms attack both l i g n i n and the polysacchar-ides, but not necessarily at the same rate. Other workers divide the 10 organisms into three types: destructive or brown rot, corrosive or inter-mediate rot, and white rot (k0,62). In this separation 'white rot' means that the fungi are able to decompose only the li g n i n component of wood, and the carbohydrates remain unattacked. In the opinion of Lyr and Ziegler (k9), however, the microorganisms originally developed the a b i l i t y to destroy cellulose, which was developed by plants e a r l i e r in time than was l i g n i n . Because l i g n i n appeared only i n the higher plants, the a b i l i t y to destroy lignin also appears only among the higher fungi (Basidiomycetes). This class of fungi i s therefore able to destroy both cellulose and l i g n i n . This means no fungi destroy lignin alone and leave the cellulose unattacked, or i n other words, no true white rot exists. Recently attention has been turned to another type of rot, to the so-called soft-rot. Soft-rot i s caused by certain species of micro fungi (Fungi Imperfecti and Ascomycetes) (52). Chemical analyses of wood decayed by Chaetomium globosum, which i s one of the soft-rot fungi, and has been most extensively studied, showed a vir t u a l depletion of carbohydrates, while the lign i n , although i t decreased steadily, was not markedly attacked. In this respect the wood was similar to that attacked by a brown-rot fungus, but instead of an increase, a slow but steady decrease of alkaline so l u b i l i t y was observed, which i s a characteristic of the white-rotting organisms.(26). Biochemical and enzymatic processes have been studied by many workers (30,32,33,kk,k9,5l,6l) i n order to get a clear picture of the de-struction of wood. The f i n a l description i s yet far from complete. No doubt i t would be attractive to form a quotient from the value of the destruction of lignin and cellulose, for characterizing the type of 11 destruction for individual fungi. Among others, Kawase (kO) used the ratio of holo-cellulose to lignin as an indicator of the type of rot. He divided fifty-one species of fungi into three types: brown rot, where the ratio of the most rotted wood i s below onej white rot, where the ratio i s high (in his study sometimes i n f i n i t e ) ; and intermediate type, where the ratio i s not far from that of sound wood. . According to these ratios he found four white-rotting fungi, twelve brown-rotting fungi, and the remaining thirty-four species belonged to the intermediate type. Lyr and Ziegler (49), investigating two white-rotting fungi (Phellinus igniarius Quel, and Collybia velutipes (Urt.), formed a quotient from the values of the percentage of the decomposed lignin and cellulose. These quotients were so changed during the course of decay that G_j_ velutipes displayed a tendency i n advanced decay to behave as a brown-rotting organism. Therefore, i n consequence of the great v a r i a b i l i t y of such a quotient, i t can be used only as an indication of the fungus, but not for separation of the type of rot. As stated, most studies on decayed wood are concerned with com-paratively advanced decay. I t i s very l i k e l y that the incipient and inter-mediate stages of decay are more important from the industrial point of view. Although decayed wood at an early stage i s widely used for making pulp, only a few a r t i c l e s are concerned with incipient decay. Scheffer (58) followed the decomposition of red-gum sap wood by Polyp orus versicolor, starting at five-per-cent decrease i n specific gravity. Cold-water solubles and hot-water solubles were unchanged, and the one-per-cent a l k a l i solu-b i l i t y slightly increased with the advance of decay. The cellulose content 12 on the basis of decayed wood remained unchanged up to thirty-two per cent weight loss, which was the highest i n his experiment. Kawase (kO) analyzed discolored wood decayed by several species of fungi, and followed the changes of the components of several wood species through the so-called more decayed and most decayed woods. MATERIALS AND PROCEDURE 13 This experiment was planned to analyze chemically alder wood decayed by the t e s t fungi to one, four and seven per cent weight l o s s . F o r t e s t organisms, Fomes p i n i c o l a (Schwartz ex F r i e s ) Cooke as a bro-wn-r o t t i n g fungus, and Stereum hirsutum F r . , Polyporus v e r s i c o l o r L. ex F r i e s , and a species of Hymenochaete as white-rotting organisms, were selected from the common decay fungi o f red a l d e r i n B r i t i s h Columbia. The l a s t fungus was rejected because i t was impossible to get uniform m y c e l i a l growth i n the incubation chambers. Rectangular glass tanks measuring 2.5 x k inches a t the bottom and 2 inches i n height were used as incubation chambers. Malt agar (3.5 per cent) was poured a f t e r s t e r i l i z a t i o n of the dishes and the malt agar solution at 110°C. Every d i s h was inoculated at eight p o i n t s with a small piece of a c t i v e l y growing c u l t u r e . When the fungi covered the malt agar completely, four g l a s s c a p i l l a r y tubes were put on the mat of mycelium, p a r a l l e l to the longer side of the d i s h , to keep the wood sections above the mat. A flat-bottomed glass tube was sealed to the side of each incubation di s h as a water rese r v o i r ( F i g . 3 ) . A i r - d r y wood was cut on a s l i d i n g microtome, i n the r a d i a l plane, i n t o pieces 10 x 30 x 0.3 mm. Specimens were t o be analyzed chemically at three stages of decay f o r each of the three t e s t f u n g i . About twelve grams of wood were needed f o r the chemical analyses. By considering the expected weight l o s s , the o r i g i n a l moisture content of the wood, and the amount of wood needed f o r the moisture content determinations before and a f t e r the 14 decay period, i t was c a l c u l a t e d that f i f t e e n grams of wood would be required. T h i s amount was prepared f o r each of the nine experimental samples and f o r the four c o n t r o l determinations. About 310 microtome sections made up the required f i f t e e n grams. The samples were a i r - d r i e d , the moisture contents were determined on twenty pieces from each sample, and the oven-dry weight of every sample was c a l c u l a t e d . The wood sections, a f t e r s t e r i l i z a t i o n f o r f i v e seconds i n b o i l i n g water, were placed i n the decay chambers having three-week-old cultures, onto the gl a s s c a p i l l a r y tubes. The s t e r i l i z a t i o n a l s o moistened the o r i g i n a l l y a i r - d r i e d wood sections (3,18,58). Each decay chamber contained, from eighteen to twenty-one wood sections. The samples were to be analyzed chemically at one, four and seven per cent weight l o s s . In order to f i n d the required time t o reach these weight l o s s e s , a preliminary experiment with numbered wood, sections was set up about three weeks i n advance of the main experiment. These wood sections had been p r e v i o u s l y used f o r the moisture-content determinations of the samples, and t h e i r i n d i v i d u a l oven-dry weights were known. Pieces were taken p e r i o d i c a l l y and t h e i r weight l o s s determined by drying them at 105°C. Using these data, curves were c a l c u l a t e d between time and weight l o s s , and the time was i n t e r p o l a t e d f o r one, four and seven per cent weight l o s s . At the c a l c u l a t e d date, the samples f o r chemical analyses were taken from the decay chambers and a i r - d r i e d . . Ai r - d r y i n g stopped the decay, gave uniform moisture content and made i t p o s s i b l e to grind the samples i n a Wiley m i l l so as to pass a 20-enesh screen. As stated, nine decayed and four c o n t r o l samples were analyzed. Of the four c o n t r o l samples, one was ground t o pass a 20-mesh and another t o pass a kO-mesh screen without 15 any s p e c i a l treatment. These were t o determine the e f f e c t of the p a r t i c l e s i z e on the chemical analyses. The t h i r d sample was analyzed a f t e r s t e r i l -i z a t i o n , while the fourth one was stored i n the wet and s t e r i l e condition i n an uninoculated decay chamber f o r f o r t y - e i g h t days. Both samples were ground to pass a 20-mesh screen. A l l samples were analyzed f o r cold-water s o l u b i l i t y , hot-water s o l u b i l i t y , methanol-benzene (1:1) s o l u b i l i t y , one-per-cent sodium hydroxide s o l u b i l i t y , l i g n i n content, h o l o - c e l l u l o s e and alpha-eellulose content (Tables 1 and 2). For the analyses each sample was di v i d e d i n t o three It-gram p a r t s . Gold-water s o l u b i l i t y was determined on the f i r s t four grams of wood-meal, followed by the h o l o - c e l l u l o s e and alpha - c e l l u l o s e determin-ati o n s j on the second part hot-water s o l u b i l i t y and one-per-cent sodium hydroxide s o l u b i l i t y were determined. The t h i r d p art was used to determine methanol-benzene s o l u b i l i t y and l i g n i n content. Cold-water solubles were determined by extracting four grams o f wood-meal with 300 ml. of cold water f or Lj.8 hours at room temperature, s t i r r i n g o c c a s i o n a l l y . The wood was then f i l t e r e d and the f i l t r a t e con-centrated under vacuum, d r i e d a t 102°C. and weighed (5l,6o). The same sample was used a f t e r a i r - d r y i n g f o r h o l o - c e l l u l o s e and alp h a - c e l l u l o s e determina-t i o n s , using a modified version of the c h l o r i t e technique (U2). The method i s as follows: from the a i r - d r i e d wood-meal 0.7 i 0.05 grams was weighed i n t o a 50-ml, Erlenmeyer f l a s k j to t h i s was added 10 ml. of b u f f e r s o l u t i o n (con-t a i n i n g 60 ml. a c e t i c acid and 20 g. sodium hydroxide per l i t e r ) , and one ml. sodium c h l o r i t e solution (containing 200 g. sodium c h l o r i t e per l i t e r ) . The f l a s k s were l o o s e l y stoppered with weighted glass stoppers, placed i n a 16 water bath regulated to 70°C. and swirled each 30 minutes. After 3/k hour, l l / 2 hours and 2 l / 2 hours, an additional one ml. of sodium chlorite solution was added. After four hours cooking, 15 ml. of cold d i s t i l l e d water was added. The contents of the flask were then transferred to a weighed 15>-ml. capacity, coarse fritted-glass crucible and f i l t e r e d . F o l -lowing t h i s i t was washed with 100 ml. of one per cent acetic acid solution, and twice with $ ml. acetone, and air-dried. The holo-cellulose was deter-mined i n t r i p l i c a t e . One of the air-dried holo-c ellulo se samples was oven-dried for moisture-content determination, and -the other two were used for alpha-cellulose determination. The holo-cellulose i n the crucible was extracted with two 3-ml. portions of 17.5 per cent sodium hydroxide solution, at room temperature, for kO minutes. After the extraction the alpha-cellulose was washed with 60 ml. of d i s t i l l e d water, f? ml. of ten per cent acetic acid, followed by another washing with 60 ml. d i s t i l l e d water, and f i n a l l y with two portions of 10 ml. acetone. The alpha-cellulose was oven-dried at 102°C. and weighed. Lignin was determined on the second k-gram sample i n t r i p l i c a t e , using a method described by Jayme, Knolle and Rapp (39)• According to the description of this method, a previous methanol-benzene extraction for six hours i n Soxblet apparatus i s needed j the extract of the whole li=-gram: : . sample i s then concentrated under vacuum, and dried at 102°C. The th i r d sample of k grams of wood-meal was used to determine the amount of hot-water-solubles and one-per-cent sodium hydroxide solubles. The hot water extraction was carried out by using 100 ml. d i s t i l l e d water for three hours at 92°G. The wood-meal was fil t e r e d , washed with hot 17 water and the f i l t r a t e concentrated under vacuum, and d r i e d a t 102°G. The wood-meal was d r i e d a t room temperature and used f o r the determination o f the one-per-cent sodium hydroxide s o l u b i l i t y by extracting the wood-meal f o r one hour at 92°C. with 100 ml. o f s o l u t i o n . The extracted wood was washed with hot water, followed by 50 ml. of ten-per-cent acetic a c i d solu-t i o n , and again with hot water. The wood-meal was oven-dried at 102°C. and weighed. RESULTS 18 From the values of weight l o s s e s of the i n d i v i d u a l , numbered microtome sections i n the preliminary experiment, the incubation periods f o r the r e q u i r e d weight losses were calculated s t a t i s t i c a l l y f o r S, hirsutum and P , v e r s i c o l o r . Curves (Figures k and 5) were used i n t h i s c a l c u l a t i o n because, t h e o r e t i c a l l y , the weight gain during the very e a r l y stages of decay makes second-degree r e l a t i o n s , instead of l i n e a r , between weight l o s s and incubation p e r i o d . For the c a l c u l a t i o n only that portion o f the data (Tables 3 and k) was used which i n v o l v e d samples most l i k e l y to have progress of decay s i m i l a r to t h a t of the samples f o r chemical a n a l y s i s . The growth of F. p i n i c o l a was more vigorous on malt agar i n the dishes prepared f o r the samples f o r chemical analyses than i n the dishes of the wood sections f o r the p r e l i m i n a r y experiments. Therefore, no equation was c a l c u l a t e d f o r t h i s fungus. The c a l c u l a t e d periods were too long f o r Pj_ v e r s i c o l o r . According to the equation, nineteen days are needed f o r one-per-cent weight l o s s , t hirty-one days f o r four-per-cent weight l o s s and t h i r t y - n i n e days f o r seven-per-cent weight l o s s . The samples f o r the chemical analyses resulted i n weight losses o f 0 . 6 , 7.1 and 21.1 per cent r e s p e c t i v e l y . A l l three samples decayed by S. hirsutum had weight gain instead of weight l o s s i n the p e r i o d based on the p r e v i o u s l y c a l c u l a t e d equation. Results of the analyses of a l l thirteen samples described i n the previous section are shown i n Tables 1 and 2. The data i n Table 1 are c a l -c u lated on the b a s i s of the weight of decayed wood. Table 2 contains the 19 data calculated on the basis of weight of the original sound wood, but only for those samples which had weight losses. 20 DISCUSSION This experiment was i n i t i a t e d i n order to study i n c i p i e n t decay e For such a study a uniform fungal attack i n the whole sample i s e s s e n t i a l . Red a l d e r i s a suitable wood species i n which to get uniform decay because i t i s not decay r e s i s t a n t , and i t s d u r a b i l i t y , therefore, does not change according to the p o s i t i o n of the samples i n the tree ( l l i , 3 7 , 3 8 ) . The wood to be exposed to decay was prepared by c u t t i n g 0,3-mm.-thick microtome sections. This t h i n section was selected to produce a natural condition f o r the t e s t f u n g i . Wood-meal was not used because i t has unnatural phys-i c a l , physicochemical and a i r conditions (35) , In l a r g e wood, blocks the fungus needs time to grow from the surface i n t o the wood, and therefore the extent of decay gra d u a l l y decreases from the surface inward (29), Probably the decay was uniform i n any one microtome s e c t i o n , but f o r the chemical analyses about 300 pieces were needed, and. these were incubated i n sixteen Incubation chambers. Only those chambers i n which the t e s t fungi grew vigorously were used, to decay wood. But even with t h i s pre-caution, the vigour of the fungi and t h e i r a b i l i t y to attack wood, varied from d i s h to d i s h , and the re s u l t a n t decay was not uniform. To prepare the chambers for one 15-gram sample, about 25 incubation chambers were inoculated i n about 200 places. In so many i n o c u l a the. mycelium of the fungus had. d i f f e r e n t v i t a l i t y . The g l a s s dishes, used as incubation chambers, were not suitable to c u l t u r e f u n g i . In many cases the cultures i n the dishes, p a r t i c u l a r l y i n the longer incubation p e r i o d , d r i e d i n s p i t e o f the presence of the small water r e s e r v o i r s . P. v e r s i c o l o r appeared the l e a s t s e n s i t i v e to the low 21 moisture content. I f i t started to d e t e r i o r a t e the wood, the humidity o f the dishes increased, and they were self-conditioned f o r fu r t h e r fungal growth. S_j_ hirsutum appeared to be the most s e n s i t i v e to the moisture content of the wood. For the determination of the stage o f decay, the weight losses of the samples were measured. In a l l cases with S. hirsutum, and i n two cases with F^ p i n i c o l a , the r e s u l t s showed weight gains i n s t e a d of weight l o s s e s . Hartley (36) reported weight gain, and according to him the weight gains were small, but they appeared too frequently to be only experimental error . Kawase (hO), i n v e s t i g a t i n g discolored wood, got weight gains up t o 11.6 per cent. This weight gain probably can be explained by the weight of the fungal mycelium, by the weight gain produced by the hydrolysis of the wood polysaccharides, and the adsorption of the components of the malt agar. These are only hypotheses, and none of them were proved, to p l a y any p a r t i n the increase of the weight of wood. The values of the chemical analyses show that c o l d - and hot-water s o l u b i l i t y increased most at the e a r l i e s t stage o f decay caused b y both white- and brown-rotting organisms. C e l l u l o s e and l i g n i n content were not changed by any of the t e s t f u n g i . This i s expected i n the wood attacked by p i n i c o l a and S_j_ hirsutum, because o f t h e i r very i n c i p i e n t stage. P.  v e r s i c o l o r destroyed the c e l l u l o s e and l i g n i n at about the same rate as they are present i n the wood., and therefore t h e i r percentage d i d not change on the b a s i s of the weight o f decayed wood. Scheffer (58) found that P. v e r s i -c o l o r destroyed red-gum-rwood i n a s i m i l a r way, but i n h i s experiment the A amount of water solubles remained the same throughout the d i f f e r e n t stages 22 of decay. Campbell's study with Pj_ versicolor on beech and ash wood (10) resulted i n an increase i n water solubles. His experiment with hirsutum on oak sapwpod, however, i s not comparable with the result of this experi-ment, because he analyzed the wood after thirteen months incubation period with a weight loss of about twenty per cent. Bray's study with Fj_ pinicola also had too high weight losses (from nine per cent up) in comparison with the weight loss caused by t h i s fungus i n the present experiment. CONCLUSIONS 23 Microtome sections, to get uniform decay in wood, are u s e f u l only i n experiment, where small amounts of wood are needed. Where a l a r g e number o f sections are required, as was the case i n t h i s experiment, i t i s impossible, under the conditions used, to produce uniformly vigorous cultures t h a t w i l l decay many wood pie c e s . Large dishes, used as incubation chambers, need very c a r e f u l handling to produce a favorable environment f o r the fungi. The dishes used, having l o o s e l y f i t t i n g l i d s , were not found to be suitable f o r t h i s purpose. At a very e a r l y stage of decay the fungi caused weight gain instead of weight l o s s . In recent l i t e r a t u r e , weight gains of more than one per cent have been r a r e l y reported, but they were w e l l over t h i s value i n t h i s experiment. Incipient decay caused, greater e f f e c t s on c o l d - and hot-water s o l u b i l i t y than on any other components determined of red alder, which was probably more a f f e c t e d by brown-rotting organism than by white-rotting fungi. 2h SUMMARY The chemical p r o p e r t i e s of red alder wood were studied a f t e r exposure t o the t e s t fungi, v e r s i c o l o r , F. p i n i c o l a , and S^ hirsutum. The experiment was designed to permit analyses at one, four and seven per cent l e v e l s of weight l o s s , A study was made of the c o l d - and hot-water s o l u b i l i t y , methanol-benzene ( 1 : 1 ) s o l u b i l i t y , one-per-cent sodium hydroxide s o l u b i l i t y , l i g n i n content, and holo- and alpha-cellulose content. In order to secure uniform decay, the wood was cut on a s l i d i n g microtome, i n the r a d i a l plane, i n t o pieces 1 0 x 3 0 x 0 , 3 mm. About 3 1 0 of these microtome sections were required f o r each stage of decay and t e s t fungi. .Glass incubation chambers containing malt agar medium were used f o r cu l t u r e s , and each chamber had from eighteen to twenty-one wood sections. Therefore, f o r the 3 1 0 wood pieces, sixteen dishes were needed. The vigour of the t e s t fungi to attack wood, i n s p i t e of a l l precautions, was d i f f e r e n t from d i s h to d i s h , and the production of uniformly decayed samples was not achieved. Up to about seven-per-cent weight gains were found i n samples decayed by S^ hirsutum and F. p i n i c o l a , but P. v e r s i c o l o r destroyed the wood r a p i d l y , and i n 3 9 days caused a weight l o s s of twenty-one per cent. A l l the fungi caused a rapid, increase i n c o l d - and hot-water s o l u b i l i t y . The increase of one-per-cent sodium hydroxide s o l u b i l i t y was most rapid i n samples exposed to F_. p i n i c o l a , but the change was slower than observed f o r the water s o l u b l e s . The det e r i o r a t i o n of c e l l u l o s e and l i g n i n appeared i n samples decayed by P_. v e r s i c o l o r , but t h e i r d e t e r i o r a -t i o n by the other two fungi was within the l i m i t of experimental e r r o r . LITERATURE Ainsworth, J.H. i960. 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U3(7)t 609-611. B r i t i s h Columbia. Department of Lands and Forests. 1957. Continu-ous forest inventory of Br i t i s h Columbia, I n i t i a l phase. B r i t i s h Columbia Department of Lands and Forests. Campbell, W.G. 1930. The chemistry of the white rots of wood. I. The effect on wood substance of Polyporus versicolor (L.) Fr. Biochem. J. 2U: 1235-12k3. . 1931. The chemistry of the white rots of wood. I I . The effect on wood substance of Armillaria mellea Fr., Polyporus  hispidus Fr. and Stereum hirsutum Fr. Biochem. J. 2£: 2023-2027. . 1932. The chemistry of the white rots of wood. I I I . The effect on wood substance of Ganoderma applanaturn (Pers.) Pat., Fomes fomentardus (Linn.) Fr., Polyporus adustus (Willd.) Fr., Pleurotus ostreatus (Jacq.) Fr., Armillaria mellea (Vahl.) Fr., Trametes~pini (Brot.) Fr., and Polystictus abietinus (Dicks.) Er. Biochem. J . 26(2): 1829-1838. 26 j ^ , and S.A. Bryant, 19kO. A chemical study of the bearing of decay by Pheilinus crypt arum Karst, and other fungi on the destruction of wood by the death-watch beetle (Xestobium rufovillosum DeG.). Biochem. J . 3k( 10-11): 3it0k-lklk. Canada, Forest Products Laboratories. 1951. Canadian woods, their properties and uses. Forestry Branch, Ottawa. 36? pp. Casey, J.P. i 9 6 0 . Pulp and paper chemistry and chemical technology. Vol. 1. 2nd Bd. Interscience Publ. Inc., New York. 580 pp. Cartwright, K. St. G., W.G. Campbell, and F.H. Armstrong. 1936. In-fluence of fungal decay on the properties of timber. I. The effect of progressive decay by Pplyporus hispidus Fr. on the strength of English ash, FraxJnus hispidus L. Set. B. Proc. Roy. Soc. London. 120, 76-851 - , and W.P.K. Findlay. 1958. Decay of timber and i t s pre-vention. 2nd Ed. Her Majesty's Stationery Office, London. 332 pp. Chapman, A.D. 1933. Effect of steam sterilization on susceptibility of wood to blue staining and wood destroying fungi. J. Agr. Res. 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Der Lignin - und der Cellulose - Abbau des Holzes, zwei verchiedene Zersetzungsprozesse durch holz-bewohnende Fadenpilze. Berichte d. D. Chem. Gesellschaft. 60(1): 225-232. Ference, G.M., and T.L. G i l l e s . 1956. The deterioration of straw-pil e d pulpwood. TAPPI. 39(6): U06-U15. Findlay, W.P.K. 1953. Influence of sample size on decay rate of wood i n culture. Timber Technology and Machine Woodworking. 6l(2l66): 160-162. Fischer, G. 1953. Untersuchung Uber den biolbgischen Abbau des Lignins durch Microorganismen. Arch. f. Microbiol. 18: 397-l}.2l4.. F r i t z , C.W. 195U. Decay in poplar pulpwood in storage. Can. J. Bot. 32(4): 799-817. Garren, K.H. 1 9 3 8 . Studies on Polyporus abietinus. I. The enzyme-producing a b i l i t y of the fungus, rPhytopathology. 2 8 : 8 3 9 - 8 U 5 . . 1 9 3 8 . Studies on Polyporus abietinus. I I . The u t i l i z a -tion of cellulose and lignin by the fungus. Phytopathology. 2 8 : 875-878. *~ Glennie, D.W., and H. Schwartz. 1950. Review of the literature on decay i n pulpwood, i t s measurement and i t s effect on wood properties. Dept. Resources and Development. Canada. For. Prod. Lab. Mimeo 0-153. 22 pp. Haden-Guest, S., J.K. Wright, and E.M. Teclaff. 1956. A world geography of forest resources. The Ronald Press Co., New York. 736 pp. Hartley, C. 1958. Evaluation of wood decay i n experimental work. U.S.D.A. For. Prod. Lab. Rept. No. 2119. 57 pp. Hawley, L.F., L.C. Fleck, and CA. Richards. 192U. The relation be-tween durability and chemical composition i n wood. U.S.D.A. For. Prod. Lab. 11 pp. Hunt, G.M. 19Ul. Factors that influence the decay of untreated wood in service and comparative decay resistance of different species. U.S.D.A. For. Prod. Lab. Rept. No. R68. 5 pp. Jayme, G., H. Knolle, and G. Rapp. 1958. Entwicklung und endgultige 28 Fassung der Ligninbestimmungs Methcde nach Jayme - Knolle. Das Papier. 12(17/18): k6k-k67. Kawase, K. 1958. Chemical study of decayed wood. Res. Bull. Exp. For. Hokkaido Univ. 19_(2): 1-330. Kennedy, R.W. 1958. Strength retention in wood decayed to small weight losses. For. Prod. J. 8(10): 308-31k. , and J.M. Jaworsky. I960. Variation in cellulose content of Douglas f i r . TAPPI. _ 3(l): 25-27. Kramer, P.J., and T.T, Kozlowski. i960. Physiology of trees. McGraw-Hill Book Co., Inc., New York, Toronto, London. 6k2 pp. Laatsch, W. 195k. Dynamik der Mitteleuropaischen Mineralboden. Verl. Theodor Steinkopff, Dresden und Leipzig. 277 pp. Lindgren, R.M. 1953. Deterioration losses in stored southern pine pulpwood. TAPPI. 36(6): 260-26U. _. 1953. An overall look at wood deterioration. U.S.D.A. For. Prod. Lab. Rept. No. I966. 5 pp. , and E.C.O. Erickson. 1957. Decay and toughness losses in southern pine infected by Peniophora. For. Prod. J. 7(6); 201-20L. , and E. Wright. 195U. Increased absorptiveness of molded Douglas f i r posts. J. For. Prod. Res. Soc. k(k): l62-l6k. Lyr, H., and H. Ziegler. 1959. 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Use of decayed wood in bleached sulphite pulp. Reprint from Paper Trade J. 5 pp. Sandermann, ¥. 1956. Grundlagen der Chemie und chemischen Technologie des Holzes. Akademisch Veragsgesellschaft. Geest und Portig K.-G. Leipzig. lr?8 pp. Scheffer, T.C. 1936. Progressive effects of Polyporus versicolor on the physical and chemical properties of red gum sapwood. U.S.D.A. Techn. Bull. No. 527 . 46 pp. , and R.M. Lindgren. 1940. Stains of sapwood and sapwood products and their control. U.S.D.A. Techn. Bull. No. 7 l U . 12k pp. , T.R.C. Wilson, R.F. Luxford, and C. Hartley. 19U1. The effect of certain heart rot fungi on the specific gravity and strength of Sitka spruce and Douglas f i r . U.S.D.A. Techn. Bull. No. 779. 2h pp. Schmitz, H. 1921 . Studies in wood decay. II. Enzyme action in Poly- porus volvatus Peck, and Fomes igniarius (L.) Gillet. J. General Physiology. 3 ( 6 ) : 795-805"; Sheridan, T.G. 1958. A mill survey of wood deterioration and i t s effect on pulp yield and quality. Pulp and Paper Mag. Canada. 5 9 ( 3 ) : 228-235. Snedecor, G.W., and W.G. Cochran. 1957. Statistical methods. 5 t h Ed. The Iowa State College Press, Ames, Iowa. 534 pp. Suolahti, 0 . , and A. Wallen. 1958. Der Einfluss der Nasslagerung auf das Wasseraufnahmevermogen der Kiefernsplintholzes. Holz Roh-u. Werkstoff. l 6 ( l ) : 8 - 1 7 . United States. Department of Agriculture, i 9 6 0 . Red alder. United States Forest Service. 1* pp. Wellwood, R.W. 1956. The manufacture and uses of hardwoods in British Columbia. A paper presented before the Northwest Hardwood Assoc-iation, Vancouver, B.C. U pp. 30 Wheeler, P.R. I960. Hardwood use from forester's viewpoint. Pulp and Paper. 3h(S): 111. Wilson, J.W., H. Worster, and D. O'Meara. i960. The sulphate pulp-ing of some South Central British Columbia wood species. Pulp Paper Mag. Canada. 6l(l): 113-118. Wise, L.E., and E.C. Jahn. 195>2. Wood chemistry. Vols. I and. II. Reinhold Publ. Corp., New York. 13U3 pp. Wright, E. 195>4. A preliminary study of the deterioration of alder and Douglas f i r chips in outdoor piles. U.S. For. Serv. Pacific Northwest For. Expt. .St. Res. Note. No. 99. E> pp. APPENDIX I (Tables) Table I Percentage of cold-water, hot-water, methanol-benzene and al k a l i solubles, Holo- andc<-cellulose and lignin in sound and decayed alder wood samples. (Percentage expressed on the basis of the weight of decayed wood). Control (20 mesh) Control (HO mesh) Control (sterilized) Control ( s t e r i l i z e d and stored) Weight Loss Cold-water Solubles Hot-water Solubles Methanol-benzene Solubility 1 per cent NaOH Solubility Holo-c ellulo se (X -cellulose Lignin Sample No. Sample No. Sample No. 1 2 3 Av. 1 2 Av. 1 2 3 Av. Per cent mm 0 0.85 0.58 0.1*8 0.38 1.10 1.10 0.97 0.85 1.23 1.19 1.18 0.96 12 .1* 10.3 13.3 12.6 83.2 82.1 81.1 85.2 82.5 81.3 80.9 83.3 83.3 80.lt 80.9 8U.5 83.0 81.2 81.0 8U.2 1*7.3 1*5.5 1*6.1 1*6.9 1*7.8 1*5.1 1*5.5 1*7.6 1*7.5 1*5.3 1*5.8 1*7.2 26.6 29.0 27.3 28.0 27.5 29.0 26.1* 28.9 28.2 29.1 26.9 28.1* 27.1* 29.0 26.8 28.1* Stereum hirsutum Stereum hirsutum Stereum hirsutum -0.60 -7.1*1* 1.56 1.25 # 2.1*5 2.27 1.85 1.1*5 1.30 1.38 11.6 12.8 13.2 81.7 83.1 8k.lt 81.9 82.3 8it.2 82.5 83.2 8it.l 82.1 82.9 81t.2 1*6.5 1*6.5 U7.9 1*7.7 1*6.5 1*8.9 1*7.1 1*6.5 1*8.1 28.5 26.5 27.7 29.7 28.1 28.6 30.0 28.5 28.0 29.1* 27.7 28.1 Polyporus versicolor Polyporus versicolor Polyporus versicolor 0.60 7.1U 21.09 1.1*3 5.38 8.1*5 2.1+1 5.89 8.1*5 0.97 2.67 U.06 7.2 ll*.9 15.0 81.2 81.1* 83.7 80.8 82.6 83.1 79.0 81.5 83.9 80.U 81.8 83.6 1*5.5 1*6.7 1*6.3 1*5.5 1*6.0 1*5.7 1*5.5 1*6.3 1*6.0 27.5 25.9 26.7 27.1* 25.6 25.7 26.5 25.3 2i*.9 27.2 25.6 25.9 Fomes pinicola Fomes pinicola Fomes pinicola -5.23 -1*.U5 2.37 5.0k I*. 72 5.65 6.96 6.69 6.1*8 U.93 1*. 73 3.66 U*.o 15.0 13.8 82.5 82.6 82.3 82.8 82.7 82.2 81.3 80.0 82.8 82.3 81.8 82.lt 1*6.1 1*6.3 1*5.9 1*5.6 1*5.5 1*5.7 1*5.9 1*5.9 1*5.8 27.5 27.7 27.7 26.8 28.0 27.2 28.3 28.0 26.1 27.5 27.9 27.1 *The sample was lo s t . ro Table II Percentage of cold-water, hot-water, methanol-benzene and a l k a l i solubles, lignin, holo- andc<-cellulose in alder wood. (Percentage expressed on the basis of the weight of original sound wood) Sample Weight Loss Cold-water Solubles Hot-water Solubles Methanol-benzene Solubility 1 Per cent NaOH Solubility Holo-cellulose C< -cellulose Lignin Per Cent Control (ster i l i z e d and stored) 1.72 0.37 0.8k 0.9k 12.k 82.9 k6.k 27.9 Polyporus versicolor 0.60 1.1*2 2.k0 0.97 7.2 80.0 k5.2 27.0 Polyporus versicolor 7.1k 5.02 5.50 2.k9 13.9 76.1 k3.2 23.9 Polyporus versicolor 21.09 6.98 7.38 3.35 12 .k 69.0 38.0 21.k Fomes pinicola 2.37 5.50 6.32 3.57 13.k 80.k kk.6 26.k KJ3 3k Table III Values of weight losses and incubation periods i n the preliminary experiment with versicolor Incubation period Weight losses in days per cent u - U . 6 , 1.1 7 -3.1, -0.7 9 l . U , 10 0.0, -8.6 15 -5.4, 0.0 16 O.U, -1.2 18 0.0, -1.7 20 2.U, -1.7 23 3.2, 3.5 26 io.5 30 17.8 3 6 U.7 UO 1.0 111 18.9, 8.U 52 2U.5, 26.0 56 7.5, 13.9 (Minus weight loss means weight gain) (Only part of t h i s data was used to c a l -culate the equation as shown i n F i g . U) Table IV Values of weight losses and incubation periods in the preliminary experiment with hirsutum. Incubation period Weight losses in days per cent 4 -lu6, 1 .1 5 3 . 1 6 1 . 5 7 - 0 . 7 , - 3 . 1 8 -9.7 10 0 . 0 , - 8 . 6 13 - 0 . 2 15 - 5 . 4 16 - 1 . 2 18 0 .0 20 - 1 . 7 , - 8 . 0 23 3 . 5 30 17 .8 31 - 5 . 6 36 k .7 kO 1 .0 k l 18.9 U2 2 . 0 44 -4 .6 kl -4.4 48 - 0 . 5 52 2U.5 56 13 .9 (Minus weight loss means weight gain) (Only part of this data was used to cal-culate the equation as shown in Fig. 5) APPENDIX II (Figures) PLATE I Figure 1. Average annual depletion in immature and mature forests of British Columbia. Figure 2. Comparison of annual decay loss with annual growth in the immature and mature forests of B r i t i s h Columbia, (in cubic feet per annum) PLATE I 1 081 b.llion cubic fast Figure 1 IMMATURE FOREST DECAY 9B MILLION MATURE FOREST GROWTH gg£ 1B4 MILLION DECAY 5S1MILLION Figure 2 PLATE II Figure 3. Butter dish as incubation chamber. Figure 3 39 Figure U. The calculated curve between incubation period and weight loss caused by P. versicolor. 30 -10 Figure 4. Uo Figure 5. The calculated curve between incubation per-iod and weight loss caused by S. hirsutum. 30 o>20j— c CD o Q. y = 0.007 x - 0.340 x + 0.8 8 c|0 to to o sz. 5 50 60 t i m e in d a y s -101 F i g u r e 5. o 

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