That'Ssó-*^ THE DIGESTION OF WOOD BY TEREDO NAVALIS E \, $>, ' BT WALTER H. DORE and ROBERT C. MILLER . •- / JJniversity of California Publications in. Zoology Vol. 22, No! 7, pp. 383-400, plate 18 UNIVERSITY OF CALIFORNIA PRESS BERKELEY, CALIFORNIA 1923■ UNIVERSITY OF CALIFORNIA PUBLICATIONS ; Nt>tfej-i-Thq Ûrfiyprsfitÿ'êf* California Publications are offered in exchange for the publi- cations* of learned societies and institutions, universities, and libraries. Complete lists, of all the publications of the University will be sent upon request. For sample copies, lists of publications or other information, address the MANAGER OF THE UNIVERSITY PRESS, BERKELEY, CALIFORNIA, U. S. A. All matter sent in exchange should be addressed to THE EXCHANGE DEPARTMENT, UNIVERSITY LIBRARY, BERKELEY, CALIFORNIA, U.S.A. WILLIAM WESLEY & SONS, LONDON A Agent for the series in American Archaeology and Ethnology, Botany, Geology, Physiology, and Zoology. ZOOLOGY.—O. A. Kofoid, S. J. Holmes, and J. Grinnell, Editors. This series contains the contributions from the Department of Zoology, from the Marine Laboratory of the Scripps Institution for Biological Research, at La Jolla, California, and from the California Museum of Vertebrate Zoology in Berkeley. Cited as Univ. Calif. 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Reaction of Various Plankton Animals with Reference to Their Diurnal Migrations, by Calvin O. Esterlyra Pp. 1-83. April, 1919 ...__________.85 2. The Pteropod Desmopterus pacificus (sp. nov.), by Christine Essenberg. Pp. 85-88, 2 figures in text. May, 1919__.....................__________________ .05 : 3. Studies on Giardia microti, by William C. Boeck. Pp. 85-136, plate 1, 19 figures in text. April, 1919............,...........................;_______ .60 4. A Comparison of the Life Cycle of Crithidia with that of Trypanosoma in the Invertebrate Host, by Irene McCulloch. Pp. 135-190, plates 2-6, 3 figures in text. October, 1919___:.__....„....._____________________...____.... .60 5. A Muscid Larva of the San Francisco Bay Region Which Sucks the Blood of Nestling Birds, by O. E. Plath. Pp. 191-200. February, 1919 ____________ .10 6. Binary Fission in Collodictyon iriciliatum Carter, by Robert Clinton Rhodes, Pp. 201-274, plates 7-14, 4 figures in text. December, 1919 .........______1.00 7. The Excretory System of a Stylet Cercaria, by William W. Cort. Pp. 275- 281, 1 figure in text. August, 1919Ë— --------------;. .........—....—„.—.10 8. A New Distome from Bana aurora, by William W. Cort. Pp. 283-298, 5 figures in text. November, 1919 .—_____________;7.____——....--------------- ^ .20 9. The Occurrence of a Rock-boring Isopod along the Shore of San Francisco Bay, California, by Albert L. Barrows. Pp. 299-316, plates 16-17, De- cember, 1919____.—...|E—--------.---.-------------------------------------— .25 10. A New Morphological Interpretation of the Structure of Noctiluca, and Its Bearing on the Status of the Cystoflagellata (Haeckel), by Charles A. - Kofoid. Pp. 317-334, plate 18, 2 figures in text. February, 1920 _________ .25THE DlBæTIOîBOF WOOD BY TEREDO NAVALIS BY gakLTER I»ORE AND ROBERT C. MILLERUniversity of California Publications in Zoology Vol. 22, No. 7, pp. 383-400, plate 18 Issued February 1, 1923 Ayric . - Forestry. Main LibraryTHE DIGESTION OE WOOD BY TEREDO NAVALIS BY WALTER H. DORE and ROBERT C. MILLER CONTENTS PAGE Introduction......................................................... 384 Biological considerations............................................ 384 Chemical considerations................................................ 386 Experimental.......... ..............jL.p.-............................ 387 Methods of analysis................................................. 388 Results........................................................... 390 Discussion of results.................................................. 392 Summary.............................................................. 397 Literature cited....................................................... 398 Explanation of plate................................................. 400 5 mini384 University of California Publications in Zoology INTRODUCTION Eveofeince the by Sellius in 17ffl, it has been debated p^pj^liEi^awhethe^)r not the destructive boring m(HlusKs5p%this group derive any nutriment- from wood wflh they» so effectively perfofafl^^eHips himself discussed the without arriving at a definit^co^lision. (177^ppjj5 ff.) expressed a nega^s opfnibnv^^^^^^ shared by Home (1814, pKg^),';*who examined wood particles removed from the bodies raM|||lanimals and thought them ^^BeSunchanged by the digestive fluids. Baumhauer (186fe,;pPl9) sta® unreservedly that TeMdo bores in wood, not£for nutrim^ffiirbut solely to secure a pro- ^^red dweMng. ®st recent writer^^Hh as Sigerfoos (1908, p. 218 Moll (19lS|p. 257 )J^^M Caiman (1919, p. 12), although considering plankton the principal food of the organisms, admit the possibility that some nou^shloent may be derived from the wood. Bartsch, however, in a recenOnonograph (1922) inclines to the earlier view of non-utilization of the wood particles as food, suggesting (p. 4) Bth apparent Bfti^^mp^that ‘ ‘ the sfiipwoKm may simply indulge in a partial meal of this kind to have the comfortable feeling of a copious repast.^y*^ BIOLOGICAL CONSIDERATIONS The arguments against uti^^®n of. the wood as gTEftL are ^Seral and varied, near relaffles of Temdo (PholaSffZirphaea, and Saxicavaj only, in such smma^sj as sandstone, limestone, shale, marl, and gneilsl—materialsj|which cannot be sup- ^^ffiiqo yield th^ffl any noqipshment; theselmmg|^i^ borings may gSem be found ^^^^^Brable quantities in the ^^^^^Sra^of the rock borers ¿{¿uaiman ,«^1919^p. 12), so that the K^^^^Sofwood borings in the digestive tra;e^^^^^^^^^^gffeitself no proof ojr utilization of 'vyood as food. The principal constitu^^Hof wood, cellulose and lignin, are notnrioiisl^r^isthnM^a^g^ive actionipM that they pass through the digesti’^Kffi^^Kf many animals^^ra’ely unchanged by the enzymes the;re|p^^n|H^g^tS® equipped t| take its food from the water as do other lamellibranchs; also, the inner end of a t^xiuP’s burrow is ofteo! closed off by the forma- tion of a shelly cross-partition, and the animal may live for an1923] D or e-Miller: The Digestion of Wood try Teredo navalis 385 indefinite period with no contac^^^^^Q wood whate^^RBi^^^S wood that has passed througSttm^^^^^H tract of the animal iS shown by microscopic study to have r^ineaEtss^Mila^^^^^^S (pi. fig. 1), and sanm inYesti^^^s*^^iicmim»ro have idefiiSa^M by its odor and Mfe. assßjßid by Moll writer remarks that at on tfi^S^LbEdMidigCStltOnids^möuci ^L more. In i^ly ¡ryju^ arguments^^ma^^^^^^Mat ^e moltEg^^^^® o"fnhe3a^g^o^^^^^^d mightXba ex^^^^Ez priori ^difi^^^^^&ts of Some^other jt^^^of animal^are known to^^^^^^^mment from notably the ter mites ( Banks^and 94) and ya^ns^^^^)oring beetles; among tha m^^^^^^^^^BsnSlkox^kn^^^^^^^H^mozymeswhich ^^mpon cellulose (Billard, 1914, p. 566). It would be surprising if Teredo a ood as ® afföRigüI by the wood^irtnu^Sn^singShSigl^^^^^^^^^^^^M While animal Me wood, ordinamf^nö this not until forced to cease boring by ^^^^^^^^mecbco^^^Hof the ^^^^^^^2*egards the woMjC'reten^n of^^ynhysiH proper^^ ^^^SnaMag^:hrough.;tlie B^estive tractive may refer ^ König’s observation (1913, p. l^^^^mt ^^^^^^^^^^^Holved from wood bvfmear^r^strorith^^^^mal lignin s^Blpoll^Ies the original EEv-gtrüctürCi Hen^^^^^uuld^x^ct,that the appearance of Jmä wood mi^^^^^Sv'Ould not be materially changed, even thüugnSh^cmulose, heimcelluloses, and soluble sugars Swerd em^ely remo^^2 The most potent^presumptive_evidence^^^^vor of. the digestion of woodKKiTered^^^^^m in tfflanatBical structure of the digest ^^^^xact. into an expanded anra elongat^^^^^^^Sth^^^^^^^^^^H. the wood pa^ticle^^Hllowed during boring ¿operates are accumulate® being then periodically Riptied int(mtmuntestin^^mence they pass into the anal canal and ar^^^emPthrough the dorsal siphoiM It seems improbable that so specialized an organ as the should have no significance in the economy ofEjMj animal except as a t^Kporary Borage for extra^^^Ssawdust on Jfs way the The anatomical ^iden(^>n this po^H will be treat^ by in a paper shortly to appear inTEfiisj series??.*SSS University of California Publications in CHEMICAL however, mufulBf inco^^^^^S The ^^ffqjW^^^BS^m^ntal hasBBBfjBl an ap]^^^S |roTtn^proBieh^ f^romTle-cheroBM ${£IBri3^^M to determine eriniMtal” (iMwBf^KBor tfloif'g^aienzvmP^^^^ Teredo are camtple of acting^plfn any^o^pife* ^^Mror not iH wood particles ^arefflKemically changed in them tHe digested trlBBP! ali^rnativePjmi heen^B^^^ftor my EHr'mgtonHi)21), whohasAhbwn, tlnmlh not s®mWusively as-; migh^^Hdesir®, that a cert ;i injB^Bm|BBnBj^s!^nm^tl|^. from livers of Wej$do norveg%jS^ill^ctfon *saw^^^tc| product Mucose. Since the liver ex^M^ failed to rediW Ph£^^Sj®ose, Harr^ngl on concludes either that the enz^^^^^^Bnlot pregent in concentration, or that motion on the limited to the hemicelliiloseB^liich are notdh’^more^BIsny broken faownmian pure cellulose, with the rarthefralternativef^hat “there may be other substanrssjpresent in wobu which * re to tlra ferment in question.” The nH^eHt utters won 1 18, pp. g parts of EftBigestiv^BBP'hSjL ^lijjsMare^11e tfffilprocess wood. o:*jfconabilmi tha^m, fluid map a me of h&aMlA^ngSiel i ujMl! mav^B! pouBujOur in Stfle region of ^^^^^So^Sto serve the do^^Spurp^^^^^^^KngJhlft&v^^^^B ja^^Hte boring and of preparing a part of it for ^^^^Boped that Harrington will ^mcal inveSL^ations farther^Tor^thelinuane himself hasBagfcl^^al. In t^^SeaiBne, it has seemed to A e^WifJ. results jnight amol)thifled^rom tHE geconq method^qlPh^BH ®tion mentionedHbove, nam^^^^^^^^^^^^^^^^^^paring tha chemical q& leav|s^^^caecum, during^H pa^^S through B^yatheruong intestine, especially the enlarged ^^^^^^pp^onBdiich ISmams a typhlosole. It seems extremely ^^^^®Sthat the material analyzed should ermnely throj£gnt the tract ih^normal^fasn^^Mand HSBiHefed through the excurrent Hhon. EXPERIMENTAL At the mar^Epanoratory maSffla^B^y t h e S an||1^>aH^c’^feay on the Oakland mole ffiMo^H|OiSough whosej^opergponf^rosM^t^^^^^^^BpJemS^^^^ ^^^g^Kor mnfg than a year pn^^riters have had unda|P®^Btion in Mm aria a consi^ffiblMiumber of Teredo namUs, whi^h highly great numbers|^KoMHFi921; have continued pga-ing anP 'aj^fflSm 1 l^^^^muerfcon5i®^^ro§ aquarium life! From tiM to time in considerable qua|^^^ftrom the ^BBIKlBpiralL borin^If^^H^arefully collected with a long pipette and dried. In about eight months sufficient material to permit® successful analysis, term 1 and iffl 2,^^pplm(lfall the mm particles used. The original wood in both cases was Douglas fir, consisting in No. 1 of squsE(Mbl(M^|plajlt^^H^^ bay for and later placed in the aquariumlfendn^^^Kl at the^arne it*was ouiiear Themmpjrial from the two aquaria has been kept separate and^mb^B^^®: p^^^S analyses, as below described, thus rendering y BEggefmW was^f^cbgnixed*of r>? especially as it|\yas not al^ffi^^^ossible^^BM^B^Ehj^Bjorin^lwith desirable promptness. In order taf^jj^B probability of such-changes, somqji^ffffl^orings388 Publi^igns in Zoology [Vol. 22 Series 1, sample 2 ^^^^Sallowed to remain on the bottom of the aquarium fol^^^^^rironmm^iefl^^pEMi'pipetted off. The boring^^^^mm^B^^^Bnried o,^®mht at lOO^^B in an oven, and crushed to pass a wire sieve h^^S 50 meshe^^H the linear inch. the ori^^i woou^er^remced to saw- KupM?M|ffground ffl passnmfgigh the 5ul^51^B]reyel To avoid |mgerlni^stiirc co^^m, all^H^^^^^pre&erve^Kor analysis in tightly fitting glass stoppered Methods of aSlysis ■i|ila|lpe& were made for moisture, extract, alcohol extract, adducing sugars in alcohol extract, hemi- gelluMgM cellulose, lignin, and furfur£ltei^J on ^filiation with hj^^^TOSric acid. All these determinations were made on one samplH of wood and the corresponding s^ple^<^^^^ffiafborings. On subse- Iffient samples some®; the determinations^^^^^^^BM Whenever P^sjht^dSf^rminationafo^cre macysimg^MgSj^rbutdn^q^ie instances ^Him&d. amount of available rnat eend upon singl^yhterminations. 1. Msodslu re«g-One gram was dried for*seven|l^hghof^mt, 1 MaliC. in a constant temperature electric oven. fibhe dried material from the moisture determination was «MatiimnB^hMnd incinerated over a wire gauze at ^^HJrelov redness. l^^^^^^wmgThSashfe^i^^^pipEi^^ahvdrohhlgnM acid and WhHSi^R^ed calcium estim^^^Hpr^pitation as oxalate in slightly DS0M#n(1 f itration of the precipi|®a^Bi permanganat JHThe conditions of essentially those! of McCrudden 4. Nitrogen.—Seven-tenths gram of material^was treated by modification of time ^i^fe^F^^pethoH:'^MiTJMtoes^^Bvere calculated as protein bv 6.25., |£ I Benzene €xirac#|^^ffl|gram of-dhq material\was weighed info^an alundum extraction thimble and dried for several hours, ^■ex|r|Kycr six hours ^^Bj^^^^Kpparat^Hl The benzene was evaporated off and the resi<^^Kh$&cl| at 100l^C. and Kveighe^H 6. Almh/jl —The residuBremainingBmtethe thimbl^ffiiffl the benzene „extraction was liyfflSfflHjwith 95 per-cent alcohol for s®1923] Dore-Miller: The Digestion of Wood by Teredo navalis 389 hours^ffi^^lcohoîi^Bporated off and^fté residual extract dried at weighed. 7. ReducTriltnsmiars in mcphol Wcfnict.—The alcohol extra^was digested in aboji^fRBc. of hot wat® allowed to cool, ant^^^^Read ^^^Sawlution added until no further^p]%llfntation occurred. ThB pr^apitate wasjfittered off and thHlear solutiondeleaded wit^^&|||n carinate and turerea. The filtrate was n^^^izedwvdth acetic acid, made up to 100 cRR and reducing sugars detèrmined by the Munson and Walk^^®906) method, the cuprous oxid^Kmgburnewto cupric oxfi||®nd^^®md as suclîfin the mannefflecommended by iJavis and 8. He-^BwMOses.—After the alcoholic extraction^ me thimble weragransferred to a 300 c.c. Erlenmeyer flask and ^^^^zea^by digesting on th^^^^Hbath for three hours with 1 per Eenrhv^ocmoric acid, the conditions being those proposecllTy Spoehr ■EETgl The unmæmved’Tbsiduë ’was fjTcred off ana^^^^gor the ^^^^^9and lignin determinations.l|||uj^*fi]^^^^^^fein^^|’ the Bifdr^jpzfd car^nwdr^^^R^!^ neutraUzelli w^^^BM«|ea^^^bonatS an^^li^^^ying BP®HPprecipitate and deleading^^^^Kffiiing sugars R^re|ld||j^^^H aM^^^^^Hunde^ ‘ reducing* £r|pffis in Cellulose and ligninllElKílt&MrimSIBB^^^^Elrate portions of for the removal of hemiS^^^^^H 9. Cellulose.-— One portion of the residual m^^j^l from hemi- cellulose determination was transferred to, a weighed Gooch crucible ^^^Sg a filtered disk then, determined by the chlorin^^B method¿Q«>iebeiifcand Walter, which hasg^^^S^eviously desjEiMd by^^^TOmWfDore, ‘ When ¿thSIfiB^^w^was appliS to bori^l^^BrjmPa cellulose residue contaminated with a certain amount of non-cellulose .jgTo^E^^^Eor the presence of mat^^Krafflud^^^^H^^sidue Was^^^^dland out ffij|rhe mixture E7 dig^sting^^^^gght in jjjhld 72jŒ^^^Rffiimuric jfâSàî The mixture ^^flli I The on the original ^^®^as!^^HrieŒ and weighed. ItS v^^H: was dedu^^Hfrom^Q weight of crude cellulose previously o^^^^^Hius giving the weight of 10. Lignin.—Another portHi of th^^^Mual material from the was a 300 c.c. Erlenmeyer390 University of California Publications in Zoology [Vol. 22 flask, 10 c.c. of ^ per cent sulfurBacid added, and the whole allowed to stand overnight. The mixture was then dilut^Swith 150 e.c. of wateOieated to boiling and filtered on a Gooch cmcible provideeljputh a disk oigiierc^^dj^oHmLhloth. I^feh^cesidue was washed thoroughly with hot water, dried^ a glassjstoppered weighing for s^^^^^mours at 100° O. and weig^las lignin. In^^^3 to learn whaler the nmj^^^^mtai^^Brom* the borings was actually ligninv iSm^dsubmi^^^^^ alie^^^Streatme^^Kvith chlorinhund 3 per centalium sulfite solrfflm. The characteristic color reactions of lignin with chlorinfe andjsulnS section were given. The treatments were repeat^ until thelulfite solution^&al nearly colorless Practically no residue then remained, indh?ajjjn3thatlthe analvt^^j product pHbably consisted Entirely of lignin. 11. Furfural yield on disiilration with hydrochloric pc^j!—One gram of the original material was treated by the method foSentosans described by the Association of Official Agricultural Chemists (1912 w Results In tables 1 and 2 below are given the Results of the analyses of wood and borings by these ^rethods. TABLE 1 OF WI As found., All figures in percentages of (.aiiS^BP. jjaaterial Determination Original ^HBorings ^BaWel" Borings Individual Average Individual Average Individual Moisture 4.89 4.91 ||90 7.36 7eBB 7.31 7.31 Ash 8.42 8|^fi| 8.22 24.17 24.17 80 MjBal M66 Calcium in ash 0.62 0.68 OKsI 0.66 o^a Nitrogen o.dM 0.08 OfÒS'j A fes o.tp 0.74 0.74 Protein 6.251. 0.50 2.27 4.62 l^^zenfftextraejH f 0:37 0.38Ì " 0.36 1 ojggw oSI OJA 1 (04 ¿jp.25, fijSS 0.44 BEf^nol’extract...... 1 7.90'* 7.90 1 |ÌAotr 1 8.00 7 nil 7.75 [27.03 25.60 26.57* Sugars in alcohol 1 0.33 0.32 1 pxtrflct' 1 0.44 0.32 j 0.35 p:ga 0. * 0.12 Hemicelluloses l(4.2Cjf fl[32 1 (as dextnffl^M i 5.04 5.24J 020 *»4'. 24 4.32 4 (PI 5.08 $| 64 5B BiEl.fi 11 osehi- ■ 47.08 47.52 47.30 14.79 £¿3.04 13.91 11.54 11W Lignin 26.47 26 41 26.44 36. W 36.11 32.13d Bay 4.75 4.53" 4.64 3.91 3.91 * 3.38 3.38* * C^^^^^aS|i*NaCl. ttSjjlgGinned.from Sgfiiage«,1923] Bore-Miller: The Digestion of Wood by Teredo navalis 391 TABLE 2 Analysis of Wood and Borings^ Series II As found. All figures in percentages of air-dried material Determination Original Wood Borings Individual Average Individual Average Moisture 8.47 8.65 8.56 7.49 7.15 7.32 Ash 6.90 7.08 6.99 18.83 18.87 18.85 Nitrogen 0.14 0.14 0.70 0.72 0.71 Protein (N X 6.25~) 0.87 0.87 4.38 4.50 4.44 Benzene extract f 0.46 0.39^ 0.40 0.42 \ 0.39 0.36 J 0.40 0.38 0.47 0.42 Sugars in alcohol extract.... nil nil nil nil nil nil Hemicelluloses Í 11.08 12.64) (as dextrose) \ 11.76 12.08/ 11.87 8.44 8.48 8.46 Cellulose 39.44 39.87 39.66 14.12 15.79 14.95 Lignin 23.63 22.91 23.27 38.16 37.71 37.93 Furfural foield 4.93 4.93 5.80 5.80 The purpose in determining moisture, ash, and protein was to enable us to reduce all data to a comparable basis. Since normal wood contains but small amounts of ash and protein, the amount found of these constituents may be taken as an approximate measure of the non-woody substances present in the samples. Accordingly, ash representMchiefly mineral contamination (shell fragments, salt, etc.) while protein represents animal remains. The analytical figureSfor the respective wood íümples. and their corresponding boring! are not directly comparable with one another since they are mixed with different amounts of non-woody material. In order to bring them to a common basis, the data have been recalculated to a moisture-free, ash-free, and protein-free basisB The results are given in table 3. TABLE 3 Recalculation of Data of Tables 1 and 2 Series I Borings ^Keries II Wood (1) (2) Wood Borings Moisture—plus ash, plus protein 13.62 33.80 42.59 16.42 30.61 Moisture-free, ash-free and protein- free material 86.38 66.20 57.41 83.58 69.39 Data Recalculated to Moisture-free, Ash-free, and Protein-free Basis Hemifeélluloses 6.02 6.46 9.33-» 14.23 12.19 Cellulose 54.74 .21.01 20.10 47.45 21.54 Lignin 30.60 54.55 55.97 27.84 54.66 Furfural yield 5.37 5.91 5.89 5.90 8.36392 of California Pufflacations in Zoology [Bre>22 The figures for benzene extracilumd alfflmJHxtract appear to havS no special significance. The former rej^^^^B resiifflSnnatter, the latter tannins, coloring matter^ and solub^ sugars. None of thesa ^cept the soluble sugars can^be regarded as ha|jmg§any nu^^^S valuelEor animals. The analyses show that extremely small amount^ of soluble sugars are present in wood and borings. Accordingly, neither benzene soluble nor alcohol soluble ^^^raients have been considered in the recalculations. DISCUSmON OF RESITI^M The data show that each of the samples of^em^ct matter contains a n^^HSmipifjm^tage of cellmcM and a muml highM^exffiniS^ pfglfnln than the wood from which it was d^M^si The figures for giemi^MlIoses and furfural yield are less regular an^Baot readily comparable'on the basis given. Inasmuch as it is extremely unlikely«that any gt^m^Byas synthe- sized during the passage of material through*h<^||f||^ve tract of |hemnimaM^^ increase in lignin is to be ascribed to concentration by glmogal. oiSetru 1 ose and other wood conAnneMlE^-Assumthe HSjp5||S^^^^^ro|^^^^ttmaining in the residue is the same as wag in the original wood, we percentage of the^ffllffld |ghM]i!i^^^^^HmanHgvi^^^H}f theti^e^aimilliof borings bij determining in each case the ra|^o£Ahe lignin content to thatSgi^a original wood. Thus the percentages of original woH substances in ^^^^^mes of borings are: » »Serirajl, sample 1 laL,:-. Series I, sample 2j Series II, The^H of tabl||l have been recal^r^^^Mfhat the (fonstiMenl^ of the nyji^^flare expressed, not on 100 parts of lK«lH but onS parts o^rigjmal wood, or, ijiothergwords, upon the frmmbcmof parts to the percen^^^found in the preceding; paragraph. The ffSfd^gare given in ^ble%. &60 54.5M 30. d| H.97 ¡.27.84 54.66 ^L00^H6.i%| X 100% = 54. S Moo^rso.MW23] The Digestion of Wood by Teredo navalis 301 TABLE 4 Data of Table 3 Calculated to Percentages of Original Wood Determination Series I Series II Woods Parts per *■" 100 "Bljings (1) Parts wea 56.1 Borings (2) Parts pér— 54.7 WoocFB Parts per Borings Parts per 50.9 Hemicelluloses 6.02 1^3,62 9M1 14.23 6.20 * o4.74 Bl 79 B99 47.45 10.96 Lignin 30.60 30.60 30.60 27.8Í| 27.84 Furfural . IÉÍ37 ' 3.22 5.90 4.261 MllMfhreS Basis, it ap]^^^ that during its“ passage through the animal’s digestive tract the wood has lostfa^^®80 per ofuts cellulosBand from 56 per ¿cení of^^^^miicell^^^^^^Kding from In 40 per fumirf|^^ldingBmstituents such as pentosanslmBp The sim^Rt explaK^^ffliof, this ^^Bpearance -?of JpirbohydratS m^ei^Mi^pfchatlthe emulóse and hemicelluloses of wood are partly ^IPsmd fch^vtiíql^^^^Han d probably ■> hydrcMzR to siiro^Mcarbo- hydrates which the animal ^^Hise as food. Consistent ’sl^^^^Higgestion tjiRMftffilteredo eonlllH*epfPx^ capable of hyfiSdyzing cellulose; also with the results of Billard fif9l|^j Bierry (191M and Bierry and Giaja who haMfound cellulose-splitting enzymes in other mo'llusks. The cellulose content ana lignin content of the three samples of I^Kigs6are remarkably ,'flffl|orm^^^^B,ble ¿PH Th i^áMi ^ iImBB indicate the composition of the residue which the teredo is incap of digesting. assumption lytical products, cellulose and lignin, are true to namer®1 for tn|^^^^^^)ased^^|ell^^^^^^^^^^^^^(Dore, 1919, 1920) question. In the ejected animal remains, there ^^Kibe some doubt as t^^^^^^^^^WBodmf^^^^^^Bution in acid is actually lignin and whether^^^^^^ffl weight which the chlorinated residue underwent really eelhimseP^- Inprderffl determine the HssibilH of animMf^Satre^^^^^^^a undissolved by*72 j p^BBi1^^^^^B»acid and contaminating the lignin, the following experinrent w^B performed: ^^Iral^tgfffii^ were taHi from the wood andfm^jg^^^Bsted out and re,^^^^B394 University ojMtalifornia Publications in Zoology [Vol. 22 ordea^Wgmiinate particlegth<^^^^^^H The flesh, thus freed from wooH]| matter, wasgfhen dried and pulverized. The residue w^S(M^ste^|^^^^ht in j$oId Rotifer and then diMf^S in the same nu^ffle as ihj^araxipg determina- tion. Solution of^®e,material^^BB^^EH^^^ta^^E^^ring gpo preserK^^nimallremains th me-lignin determination. In tMimse&jmhe celluloH consider able^mmui It, v was experier^^ in filtering and washing the materia ^i^MB^^^MndlcKTorination. ThisiattSamaBto'be due to the nSse^^Bf mass. On dissolving out the cellulose fro^^^^^^^Bue by '^SpeS ^mi^ulfuric acid, considerable amounts of insoluffi substanc^&ere obtained, as shown by table 5. TABLE 5 Cellulose Content of Wropÿnfi) BoRtNGs^i (Expressed in grams obtained from 1 gram of mnginal matex^™i Series lsUsarniilc of borings Series I 2d sample of POTm©» Series II MSiTOmj^tooringsJB (1) (2) (2) ïtesiàue afjfer chlorination Residue after acid digestion Bde^mose bv difference 0.2620 0,1141 0.26fjN 0.1357 0.1420 5 0266 0.2677 0.1265 0.2714 BBIEa 0.1479 1.1304 1 0.1154 oEI “1* not believed that^Me insoluble r^^ffe cdnMned*a-:^reat^Bl matter, as such matei^®p pears to lie Mrgen^^Mble ia [r2 percent acid under the conditions, emplmml. The residr^^^^^S sidered as ®^^fflngdalm^K liffh^whicltt^ff^^^^çpination, me partl^^^^TOr^Bec 1 i\'e a^^rn offfi^solloidal mass andHuU&ISa the Eact n at i on ^^Hnot coas Kacount o difficMdes. I^ffio be of boring^in ^^^^^^onsiderably m.mB protein thai8|heof b^^^S of th^^^^Rer^^Bee much smalleS yield of insoluble ma^B in the cel 11Uo^ide^^^^mtion, and thP ^^PeLfor and lignin similar other samples. for cellulose and lignin fare not aHcted by the amount of animal BéEJ ;^Ësent ml the sample. IO^an, be shown Ihah. even de^tive, possible allow^^M for *^^uamin^on of pr^^^Hioes nO^Eivalicm^yte^^neral conclusion thatThe Dige^mn of WHZ by [¡¡m^^mriavaUs 395 contain a smaller proportion of cedióse to lignin than the wood fem which they wer^^^^^ffl If tl^^^lulose figure is low, the correct value can in no case be higher than the total residue after chlorination (see talffiô) and if tirenignin is high, due to contamina- tion with animal remains, the nomfct value can in no pjs^be lower than the lignin v3fi|ÏP given minus the total protein of the sampl^ From tables 1, 2, and i^^can then calculate the following: * TSeries I Borings Series II (1) (2) Borings Mayimiim pnssihlp va hTp|it'<|r Minimum possible value for lignin 26.40 33.84 14.20 27.5T 26.95 33.49 In every'teasef the cellulose is materially less than the lignin, tlie origlnahy,#OT||he about twice+a^muel^ It raP^mjMSle,!^»)1-^^ drawwfrom Hfaé^daia any other concmsion than 'PelMd$e partly disappears during digestion. We have Bhe possibiliftithat the change in thHsompcSiion of the wood may oa^pl aftemtóie material isfl^Rted by^^S borer. The compffl|™nswi^tne first and ^Bcondj samples|j|f borinS| in Series-1 furnish*#«® experimental evi^^Sbn^^Hj poinl| Although P|®®nd^sam]lfé was in the sea water foil at ^^^MnaJBbntli sBpnger thatí^the^ first, thero^was noPy0j||pfeÍ reduction in carbohydrate constituents (see table 3). In*VÍeytÉIof remi extrem^^^^^fflHBIf wood to purely chemical!|Kction, and í$épbspre!| of known cellulose-dest roying fungi orKo^mhiaKn pi^Pj^^onable' to ascribe the jraS^apellulose and other carbohydrates to^E^mive ^Bln while in the body than to external agen'djbfÉPl Plankton® doubtless the principal |£eod of TerjRio^as^of other lamellilpgne^& and bn? account of its high proiá™Bm^at¿s1torobablv pufflMftf growth andiftpair of j^HySBMkereda which tly growing to fill the larger (BSály, reauife^^H^^lp protein fcodlip^Bflé for growth addi^Rial aMjuft ogjfood to furnish energy for its boring act^jj^il This Iwerg^lHnld bAscipplied by^haBgiteein material of* ttidtplankton, but much morer€^^patiy by carbohydrates,* inasmuch as thes^piast are complegSioxidized and there|is therefore no necessity*of getting rid of nitrogenous products. If, then, when its boring activities are griajlyl^the teredo has available a considerableygupmy of carbo- hydrMe material to furnish the necessary energy and that available i3396 University of California Zoology [HK| proportion to the amount of woroo be done, it appears to be a most natural arrangement, admirably adapted to its needs. The carbo- hydrates of theTwood therefore play an importantnTart in sfflplying Bheif^^o^wi^SinCTgy when it is most needed. It does notmppear probable that Teredo could ^^^^^^Sany long ffiuiod upon the wood alone,' bemuse of the negligibly amount of protein material. Especially during boring would sideraBuB nitrogenous matter ure" required. The need1 for a simuS taneous supply of plankton is accordingly indicated. *If the conclusion is accepted that wood tissue is partly digested and absorbed as food by Teredo, some ligiraHiP thrown upon the probable mecnanism by which toxic substances irijecteeHifi'to the timbers of marine structures protect Against teredo attack. If partial pige&tion of wood occurs, i®l clear that all substahcesTontained iS the wood must experience an intimate contact with the terepxact. et natur., 1, 1-45, 4 pis. Bierry, H. 1914. Ferment^ digestifs chez Helix pomatia. Compt. Eend. Soc. Biol, de Paris, 76, 710-71g Bierry, H., and Giaja, J. 1912. Untersuehungen über die Mannane, Galaktane und Cellulosen angrei- fende Enzyme. Biochem. Ztschr., 40, 370—389,|^ Billard, G. 1914. Note sur les ferments hydrolysant les hydrates de carbone chez 1’ Helix pomatia. Compt. Bend. Soc. Biol, de Paris, 76, 566-567. Kîalman, W. T. 1919. Marine boring animals injurious to submerged structures. Brit. Museum, Nat. Hist., Econ. Ser., 10, 35 pp., 21 figs, in text. Davis, W. A., and Daish, A. J. 1913. A study of the methods of estimation of carbohydrates especially in plant extracts. Jour. Agr. 'Sei., 5, 437-468fi| Dore, W. H. 1919. The proximate analysis of wood. Jour. Indus, and Engin. Chem., 11, 556-563.* 1920a. The determination of cellulose iif wood. Ibm., 12, 264-269. 1920&. Th© proxiiKfe analysis of coniferous woods. Ibid., 12, 476-479. HarriAton, C. B. 1£S1jj A note on thei physiology of the ship-worm (Teredo^iorvegica)k Biochem. Jour., 15, 736-741, 1 fig. in text. Home, E. 1814. Lectures on comparative anatomy (London, G. and W. Nicol), 1, 484 pp. Kofoid, C. A. 1921. The marine borers of the San Francisco Bay region. Beport on the San Francisco Bay Marine Piling Survey, 1, 23-61, 36 pis., 1 fig. in text.923] Dore-Miller: The Digestion of Wood hy Teredo navalis 399 Kofoid, C. A., and Miller, E. C. 1922. Specific status of the Teredo of San Francisco Bay. Iiid., Biological Section, 2, 81-82. König, J. 1913. Die Formelemente der Zellmembran, ihre analytische Bestimmung und technische Bedeutung. Chem. Ztg., 36, 1101. McCrudden, F. H. 1909. Determination of calcium in plant tissues. Jour. Biol. Chem., 7, 83-100. Moll, F. 1914. Die Bohrmuschel (Genus Teredo Linné). Naturw. Ztschr. f. Forst- u. Landwirtschaft, 12, 505-564, 12 figs, in text. Munson, L. S., and Walker, P. H. 1906. The unification of reducing sugar methods. Jour. Am. Chem. Soc., 28, 663-686. Nelson, T. C. 1918. On the origin, nature, and function of the crystalline style of lamellibranehs. Jour. Morph., 31, 53-111, 17 figs, in textJB Beimarus, J. A. H. 1773. Betrachtungen über die besondern Arten der thierischen Kunsttriebe (Hamburg, J. C. Bohn), 232 pp. Sellius, G. 1733. Historia naturalis Teredinis seu Xylophagi marini tubulo-eonchoidis speciatim Belgici (Trajecti ad Ehenum, Besseling), 353 pp., 2 pis. Sigerfoos, C. P. 1908. Natural history, organization, and late development of the Tere- dinidae, or shipworms. Bull. U. S. BuigFish., 27, 191-231, pis. 7-21. Spoehr, H. A. 1919. The carbohydrate economy of cacti. Publ. Carnegie Inst. Wash., 287, 79 pp.EXPLANATION OF PLATE PLATE 18 Eig. 1. Wood borings ejected by Teredo, magnified 300 diameters. Observe that in many of the particles the cell walls are visible. Eig. 2., Siphons protruding from blocks of wood containing Teredo. Photo- graphed in aquarium, natural size. The long siphons are the incurrent ones. The excurrent siphons are relatively short, usually protruding only two or three millimeters from the openings of the burrows. Note ejected wood borings around many of the openings. [400]• • o • ( » J t»Æ 2 m W '■» r » * * * > i ^ •■>••• p * • « ’«/«', • * * • » i » » * • * i * '<* » ? » ** <** % _ 4 % *•** « * « t • « • 4 9 % '*« - */ * «'*» * a * « f • » ,» * a * « * A « If « « » 4 * 1 * » t 9 4 It 9 * « #UNIVERSITY OF CALIFORNIA PUBLICATIONS— (Continued) 11. The Life Cycle of Echinostoma revolutum (Froelich), by John C. Johnson. Pp. 338-388, plates 19-25, 1 figure in text. May, 1920 .11______________ .6* 12. On Some New Myriopods Collected in India in 1916 hy C. A. Kofoid, hy Ralph V. Chamberlin. Pp. 389-402, plates 26-28. August, 1920 _______' .20 18. Demonstration of the Function of the Neuromotor Apparatus in Euplotes 5.00 VoL 24. 1. A Geographical Study of the Kangaroo Rats of California, by Joseph ^ Grinnell. Pp. 1-124, plates 1-7,24 figures in text. June, 1922 ......__________... 1.75 2. Birds and Mammals of the Stikine River Region of Northern British Columbia and Southeastern Alaska, by H. S. Swarth. Pp. 125-314, plate 8, 34 figures in text. 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