PROCEEDINGS OF THE a Biological Society of Washington Volume VI. February 8, 1890, to December 26, 1891. WASHINGTON, D. C. PRINTED FOR THE SOCIETY. 1892 I PUBLICATION COMMITTEE. CHARLES D. WALCOTT, Chairman. R. E. C. STEARNS, F. H. KNOWLTON, T. S. PALMER, F. V. COVILLE. II CONTENTS. PAGE Officers and Committees for 1891 iv Proceedings, February 8th to December 26th, 1891 v-xix Addresses and Communications : On Dynamic Influences in Evolution, by Wm. H. Dall (read March 8th, 1890) i-io Neo-Darwinism and Neo-Lamarckism, by Lester F. Ward (January 24th, 1891I 11-71 III LIST OF THE OFFICERS AND COUNCIL OF THE BIOLOGICAL SOCIETY OF WASHINGTON. Elected January io, 1S91 OFFICERS. PRESIDKNT. C. HART MKRRIAM. VICE-PRESIDENTS. C. V. RILEY, RICHARD RATHBUN, FRANK BAKER, C. D. WALCOTT. SECRETARIES. L. O. HOWARD, F. A. LUCAS. TREASURER. F. H. KNOWLTON. COUNCIL. C. HART MERRIAM, Presidetit. FRANK BAKER, RICHARD RATHBUN, T. H. BEAN, THEOBALD SMITH, WM. H. DALL,* R. E. C. STEARNS, THEO. N. GILL,* F. W. TRUE, G. BROWN GOODE,* GEORGE VASEY, L. O. HOWARD, C. D. WALCOTT. F. H. KNOWLTON, LESTER F. WARD,* ' F. A. LUC.\S, C. A. WHITE,* C. V. RILEY. STANDING COMMITTEES— 1891. Delegates to the Joint Commission of Scientific Societies of Washington. C. HART MERRIAM, LESTER F. WARD, RICHARD RATHBUN. Committee on Com m unications . RICHARD RATHBUN, Chairman. W. B. BARROWS, JOHN MURDOCH. Committee on Publications. C. D. WALCOTT, Chairman. R. E C. STEARNS, T. S. PALMER, F. H. KNOWLTON, F. V. COVILLE. Committee on Trees and Shrubs. LESTER F. WARD, Chairman. GEORGE VASEY, THEODORE HOLM, F. H. KNOWLTON, F. V. COVILLE- * Kx- Presidents of the Society. IV PROCEEDINGS. One Hundred and Fifty-fifth Meeting. Febrtiar}- 8, iSgo. The President in the chair, and thirty-eight members and guests present. The following active members were elected on recommenda- tion of the Council : T. \V. vStanton, U. vS. Geological Survey ; Dr. E. Roome, Columbian University. Dr. Frank Baker presented a communication upon An Undescribed Muscle from the Infraclavicular Region OF Man. Discussed by Drs. Merriam and Riley. Mr. C. D. Walcott presented a note upon A Xew Genus and Species of Ostracod Crustacean from the Lower Cambrian.* Dr. Cooper Curtice read a paper on The Moultings of the Cattle Tick {Ixodes bovis).!: Di.scu.ssed by Drs. Riley and Theobald Smith. Prof. Lester F. Ward spoke of Flowers that Bloo^i in the Winter Time. One Hundred and Fifty-sixth Meeting, February 22, 1890. The President in the chair, and thirty members present. Mr. F. V. Coville presented a paper on The New Arrange- ment OF Genera in the Herbarium of the Department OF Agriculture. + Discussed by Messrs. Riley and Ward. * Tenth Annual Report of the Director U. S. Geol. Survey, Pt. I, 1890, pp. 625, 626. tjournal Comp. Med. and Vet. Archives 1S90, p. 313 ; reprinted in Veterinarian 1891, p. 680. X Bot. Gazette, xv, 1890, p. 68. VI BIOLOGICAL SOCIETY OF WASHINGTON. Dr. T. H. Bean presented Notes on Some Fishes from British Columbia.* Dr. Merriam read a paper on the Evidence of Sonoran Origin of the Flora and F'auna of the Gulf States. Discussed by Messrs. Ward, Gilbert, Coville, and Dall. One Hundred and Fifty-seventh Meeting, March 8, 1890. The President in the chair, and fort3'-one members present. Mr. B. T. Galloway read a paper giving the results of some observations on an apple disease caused by the fungus Gym- nosporanoitiDi macropus. I^iscussed by Messrs. Van Deman and Marlatt. Mr. C. L. Hopkins presented some NoTES on the Animal LIFE above the Snow Line on Mt. Shasta. t Discussed by Messrs. Mann, Riley, Howard, Diller and Hasbrouck. Dr. W. H. Dall read a paper entitled On Dynamic Influ- ences in Evolution. j: Discussed by Messrs. Coville, Mann, Fernow and Ward. One Hundred and Fifty-eighth Meeting, March 22, 1890. The President in the chair, and thirty-nine members present. Dr. C. H. vStowell was elected an active member. Dr. D. W. Prentiss read a paper entitled Change in the Color of Human Hair,§ Change in the Color of Plum- age OF Birds, and in the Fur of Mammals. || * Proceedings U. S. Nat. Mus., Vol. xii, pp. 641, 642. t Insect Life, Vol. 2, p. 355. X Proc. Biological Society, Vol. vi, pp. i-io. § Phila. Med. Times, xi, 1880-81 ; also, Therap Gazette, Detroit, Mich., 1889, viii. llJournal Amer. Med. Assoc, xiii, 1889. PROCEEDINGS. VII Prof G. B. Goode read a paper on the Colors of Fishes.* Prof. C. V. Riley spoke on the Colors of Insects. The papers were briefly discussed by Messrs. Mann, Dall, Seaman and Ward. One Hundred and Fifty-ninth Meeting. April 5, 1890. The President in the chair, and twenty-five members present. Dr. Theobald Smith read a paper entitled Some Illus- trations OF Ferments and Fermentation among Bac- teria.t Discussed by Professor Atwater. Dr. R. R. Gurley presented a paper on the American Graptolites. One Hundred and Sixtieth Meeting, April 19, 1890. The President in the chair, and fortj^-five members present. Dr. C. W. Richardson was elected an active member of the Society. Dr. Merriam presented a paper entitled Historical Re- view OF THE FaUNAL AND FlORAL DIVISIONS THAT HAVE BEEN Proposed for North America. Discussed by Dr. Dall. Prof. J. P\ James read a paper entitled Variation with Especial Reference to Certain Paleozoic Genera. Discussed b\' Professor Ward. Dr. Dall exhibited some Original Drawings of the Fur Seal and Stellers' Sea Cow.t and gave an account of them. * Trans. Am. Fisheries Society, 1890. t Centralblatt fiir Bakteriologie u Parasitenkunde, vii, 1890, p. 502. X Report U. S. Coast Survey and Geodetic Survey, 1890. viii biological society of washington. One Hundred and Sixty-first Meeting, May 3, 1S90. The President in the chair, and thirty-six members present. Mr. J. R. Edson was elected an active member. Dr. Robert Reyburn read a paper entitled The lyiFE History of Micro-organisms with its Relation to the Theory to Evolution.* Discussed by Dr. Theobald Smith, Dr. Salmon, Mr. True, Mr. Erwin vSmith, Dr. Shaeffer, Mr. Seaman and Dr. Dall. Dr. Vasey read a paper entitled A New Grass Genus. t Discussed bv Messrs. Holm, Coville and Prof. Ward. One Hundred and Sixty-second Meeting, May 17, 1890. The President iri the chair, and thirty -six members present. Mr. T. S. Palmer read a paper entitled Some Early Views OF Geographical Distribution of vSpecies. Discussed by Dr. MerriauL Mr. F. W. True exhibited a specimen of Lophiomys imhausii. Discussed by Dr. Merriam. Prof. W. H. Seaman read a paper entitled The Place of Biology in the Public Schools.! Discussed by Profs. Chickering and Ward, Dr. Baker and Messrs. Waite and Howard. One Hundred and Sixty-third Meeting, Ma}' 31, 1890. The President in the chair, and twenty-six members present. Dr. C. H. Merriam exhibited specimens of sundry new species of North American Mammals. Discussed by Dr. Gill. * American Monthly Microscopical Journal, June, 1890. t Bot. Gazette, xv, 1890, p. 106. J The American Anthropologist for October, 1891. TROCEEDINGS. IX Dr. Theodore Gill presented a paper on the Character- istics OF THE HaLOSAUROID FiSHES.''^ Dr. J. N. Rose read a paper entitled CoultercUa, A New Genus of Composit.E.I Discussed by Messrs. Seaman, \^asey and T. S. Palmer. Prof. Jos. F. James spoke on Organisms in St. Peter's Sandstone. Mr. B. P. Mann made some remarks upon the Authorship OF THE BiBEIOGRAPHY OF ECONOMIC ENTOMOLOGY, published by Department of Agriculture. Mr. Howard refuted Mr. Mann's statement. One Hundred and Sixty-fourth Meeting, October i8, 1890. The President in the chair, and twenty-eight members present. Mr. H. E. Van Deman spoke of Cultivated Fruits in THE Mountains of North Carolina.:}: Discussed by Dr. Salmon . Dr. Theo. N. Gill presented a communication on the Super Family Cyclopteroidea.§ Prof. Lester F. Ward spoke on the subject of American Triassic Flora. 1 1 One Hundred and Sixty-fifth Meeting, November i, 1890. The. President in the chair, and thirty-seven members present. Mr. Nathan Banks was elected an active member. * American Naturalist, xxiii, 1889, pp. 1015, 1016. (Pub. May, 1890.) tr. vS. Cont. Nat. Herb, i, 1890, p. 71. J Ann. Report U. S. Dept. Agriculture, 1890, pp. 410, 411. I Proc. U. S. N. M. xiii, 1890, pp. 361-376; pi. 2S-30. II Geological Society of America, iii, 1891, pp. 23-31. Abstracts in Science, Vol. xviii, Nov. 20, 1891, pp. 287, 288, and Proc. A. A. A. S., Vol. xi. X BIOLOGICAL SOCIETY OF WASHINGTON. Mr. F. V. Coville read a paper entitled Fruiting of the Ginkgo at the Department of Agriculture. Discussed by Dr. Riley, Mr. Seaman and Mr. Fernow. Dr. Marx spoke of his recent Investigations of the Poison Glands of Lathrodectus. Discussed by Drs. Riley, Dall and Theobald Smith. Prof. Jos. F. James read a paper called Fucoids and Other Problematic Organisms. Discussed by Mr. Lucas, Dr. Dall, Prof. Ward and Mr. Mann. One Hundred and Sixty-sixth Meeting, JsTovember 15, 1890. The President in the chair, and forty-three members present. Dr. Merriam presented a communication entitled Life in the Lava Beds and Canons of Snake River, Idaho, in October.* Discussed by Messrs. Walcott and Howard. Mr. Theodore Holm spoke of the VEGETATIVE Propaga- tion OP DiCENTRA CUCULLARIA.f Dr. Dall presented some Paleontoi^ogical Notes from the Northwest Coast.]: Mr. Lucas described a Foot Disease of Captive Birds. One Hundred and Sixty-seventh Meeting, November 29, 1890. The President in the chair, and forty-seven members present. The following new active members were elected : Jno M. Stedman, Merwin M. Snell and Rev. Alexis Orban. Dr. T. H. Bean presented a paper on The Death of Salmon After Spawning. § Di.scussed by Dr. Gill, Prof. Goode, Mr. Stejneger, Drs. Dall and Merriam. *(In part.) North American Fauna No. 5, July, 1891, pp. 6-7. fBull. Torrey Botanical Club, Vol. xviii, 1891, pp. r-ii, pi. cxi-cxiii. X Nautilus, Philadelphia, Vol. iv, 1S90, No. 8, pp. 87-89, December. 'i Forest and .Stream, November 27, 1890. PROCEEDINGS. XI Dr. Theobald Smith spoke Ox Species Among Bacteria. Discussed by Mr. True, Dr. Gill, Drs. Riley, Curtice. Mr. Banks. Mr. Sudworth presented a communication entitled XoTES ON Nomenclature. Discu.ssed bv Dr. Merriam. One Hundred and Sixty-eighth Meeting. December 13, 1890. The President in the chair, and twenty-five members present. Mr. A. B. Baker was elected an active member. Mr. Wm. Palmer read a paper on The Occurrence of THE Asiatic Cuckoo on the Pribylov Islands. Dis- cussed by Dr. Dall. Dr. Riley presented some New Notes on the Genus Phylloxera. Mr. True spoke on The Teeth of the Muskrat. Mr. Lucas read a paper on The Wing of Metopidius. Discussed bv INIr. True. One Hundred and Sixty'-ninth Meeting, December 27, 1890. The President in the chair, and twenty-six members present. The following active members were elected : J. M. Holzin- ger, Frederick C. Test. Dr. Cooper Curtice presented a communication entitled A Preliminary Study of Ticks in the United States. Discussed by Dr. Smith. Dr. C. Hart Merriam exhibited A New Rabbit from the Snake Plains of Idaho.* Dr. W. H. Dall read a paper entitled On the Topography of Florida w^th Reference to its Bearing on Fossil Faunas, t * {Lepus idahoensis) North American Fauna No. 5, July, 1891, pp. 76-78. t Bulletin 84, U. S. Geol. Survey. (In press.) xii biological society of washington. One Hundred and Seventieth Meeting. (Tenth Annual.) The President in the chair, and forty-two members present. The annual reports of the Secretary and Treasurer were read and accepted. The following board of oflBcers was elected for the ensuing year : President — C. Hart Merriam. Viee-Presidenis — C. V. Riley, Frank Baker, Richard Rath- bun, and C. D. Walcott. Secretaries — L. O. Howard and F. A. L^ucas. Treasurer — F. H. Knowlton. Mernbers of Council— V . W. True, T. H. Bean, R. E. C. Stearns, Theobald Smith, and Geo. Vasey. One Hundred and Seventy-first Meeting, January 26, 1891. (Eleventh Anniversary Meeting.) The eleventh anniversary meeting was held in the law lecture-room of Columbian University, January 26, 1891, in the presence of members and invited guests. The President, L^ester F. Ward, delivered his annual address on the subject Neo-Darwinism and Neo-Lamarckism.* One Hundred and Seventy-second Meeting, February 7, 1891. The President in the chair, and twenty-five members present. The President announced the following committees for the ensuing year : Joint Commission — C. Hart Merriam, Lester F. Ward, Rich- ard Rathbun. * Published in this volume. See pp. ii-71. PROCEEDINGS. XIII Comjnittee on Comnninications — Richard Rathbuii, Walter B. Barrows, John Murdoch. Co}mnittee on Publicatiojis — C. D. W'alcott, R. h^. C. Stearns. F. H. Knowlton, T. S. Pahner, F. \'. Coville. Committee on Trees and Shrubs — Lester F. Ward, Geo. Vasey, F. H. Knowlton, Th. Holm, F. V. Coville. Mr. C. D. Walcott presented a paper on the Discovery OF Vertebrate Life in Lower Silurian (Ordovician) Strata.* Discu.ssed by Dr. (.rill. Prof. Henry F. Osborn gave a Review of the Discovery OF Cretaceous Mammals. t Discus.sed bv Dr. Gill. One Hundred and Seventy-third Meeting, February 21, 1891. The President in the chair, and twelve members present. Dr. Cooper Curtice read a paper entitled Some Little Known Worms in Cattle. Discussed by Mr. Holzinger. • One Hundred and Seventy-fourth Meeting, March 7, 1891. The President in the chair, and twenty-five members present. Mr. F. A. Lucas exhibited and described .some young Hoatzins. Discussed by Dr. Dall. Mr. Lucas also drew attention to a Specimf:n of Bison LATIFRONS FROM PEACE CrEEK, FLORIDA. Dr. Bean spoke of the Fishes of Gre.yt South Bay.;}: Mr Rose spoke of A New Aster from Southern Cali- fornia.! Discussed by Mr. Holzinger and Mr. Waite. *Bull. GeoL Soc. Amer. VoL iii, 1891 {in press). See also Proc. Phila. Acad. t American Naturalist, xxv, Juh-, 1S91, pp. 295-611. See also Proc. Phila. Acad. X Nineteenth Report of the Commission of Fisheries of New York, 1891, pp. 237-281. ^Bot. Gazette, Vol. xvi, 1891, p. 113. XIV BIOLOGICAL SOCIETY OF WASHINGTON. Mr. Slid worth presented a communication on the Color AND Odor of Flowers in Attracting Insects. Dis- cussed by Messrs. Howard and Marlatt. Mr. Stedman exhibited and described a fine specimen illus- trating the Embyro of a Chick with Two Protovertebr.'E. Dr. Merriam spoke of Distribution of Animal and Vegetable Life. Discussed by Dr. Curtice and Messrs. Waite, Test and Holzinger. One Hundred and Seventy-fifth Meeting, March 21, 1891. Ex-President Ward in the chair, and twenty-eight members present. Mr. W. A. Taylor was elected an active member. Dr. Dall presented a paper on the Age of the Peace Creek Bone Beds in Florida.* Dr. Shufeldt read a communication on A Collection of Fossil Birds from the Equus Beds of Oregon, t Mr. F. A. Lucas spoke of A Point in the Anatomy of Hesperornis. Mr. F. H. Knowlton presented a communication entitled What Are Cypress Knees ? Discussed by Prof. Ward and Mr. R. L. Garner. One Hundred and Seventy-sixth Meeting. April 4, 1891. Vice-President Walcott in the chair, and thirty-two mem- bers present. Dr. Cornelius B. Boyle was elected an active member of the Society. * Bulletin 84, U. S. Geol. Survey. (In press). t To be published in the Acad, of Nat. Sciences of Phila. Abstracts . 2.SS-259. PRESIDENTIAL ADDRESS. 19- The cogency of Lamarck's reasoning, especially when we consider the time at which he wrote, is sufficiently apparent to all, but it may not be without interest to note the manner in which it struck so excellent a judge as Professor Huxlej' as late as 1876. In contrasting it with the views of Cuvier who maintained the fixity of species and their special crea- tion, Professor Huxley says: "It is impossible to read the ' Discours sur les Revolutions ' of Cuvier, and the ' Principes ' of lyamarck without being struck with the superiority of the former in sobriety of thought, precision of statement, and coolness of judgment. But it is no less impossible to consider the present state of biological science without being impressed by the circumstance that it is the conception of Lamarck which has triumphed and that of Cuvier which has been van- quished ... It is not too much to say that the facts of biology known at the present day are all consistent with and in favor of the view of species entertained b}^ Lamarck, while they are unfavorable to, if not incompatible with, that advo- cated by Cuvier."* DARWINISM. Darwin was acquainted with Lamarck's views when he wrote the Origin of Species, and notwithstanding the fact that whenever he refers to Lamarck, as he does in several of his lettersf he does so in a very disparaging wa3% he must have been greatly influenced by them, or at least by views of the same import expressed by others as well as by Lamarck, but especially those of his grandfather Erasmus Darwdn, who anticipated, rather from the standpoint of the poet and seer, the truths to which Lamarck was led by a life-long study of living things. *Am. Cycl. , Art. Species. t Life and Letters, Vol. I, p. 542, Vol. II, p. 19b. 20 BIOLOGICAL SOCIETY OF WASHINGTON. But Darwin, like most other thorough naturalists, was little satisfied with the L,amarckian theory, because it left, as all now admit, so much still unexplained, but instead of reject- ing it in toto, as most other naturalists did, he sought, and happily succeeded in finding the principle on which the remainder of the facts could be accounted for ; or, at least, the greater part of them, for it seems that however deeply we may probe the secrets of nature there will ever remain a few residual phenomena that refuse to submit to our canons. It is certainly unnecessary that I should occupy your time with any extended exposition of the law of selection, and I will content myself with the following bare definitions : Natural selection is the general law that variations are con- stantly occurring in organized beings, and that such of these variations as prove advantageous to the species are preserved through heredity and transmitted to posterity while those which are not advantageous or are disadvantageous to the species are not preserved nor transmitted ; the cause of such selection being the fact that advantageous variations tend to increase the chances that the individuals possessing them will reach the reproductive age and continue longer to reproduce, and will hence leave a larger number of offspring than those individuals which had not varied or had varied in an equivo- cal or unfavorable manner. Artificial selection is the act of man in intelligently select- ing the individuals that possess in the highest degree the particular qualities that he desires to produce as the parents of the animals or plants which he wishes to domesticate or cultivate. The eminent success obtained by man in this way is the certain proof that the qualities of the parents are trans- mitted to their offspring, and explains the efficacy of natural selection. PRESIDENTIAL ADDRESS. 21 Sexual selection is the law that one sex, usually the female, exercises a choice between the individuals of the other, whereb}' those individuals possessing the selected qual- ities stand a much greater chance to have the opportunit3^ of transmitting them to their offspring. This law explains the greater ornamentation of the males in so many species, since most such characters are peculiar to one sex and only appear at maturity. Sexual selection also checks the tendency of natural selection to extreme variation in certain directions, since the sexes are well known to prefer their opposites, which causes the offspring to occup)- a mean between the extremes. This effect is very marked in the human race, but is doubtless operative among the lower animals. As I pointed out in last year's address, sexual selection has wrought a great revolution in the relative size, strength, and beauty of the two sexes, and reversed in birds and mammals the normal law of female superiority which prevails in most of the lower departments of life. ACQUIRED CHARACTERS. It will be readily perceived from what has been said of the two great principles of transformism, the functional, as set forth by Lamarck, and the selective, as elaborated by Darwin, that the fundamental distinction between them is that in the former the transforming qualities which are to be cumulativelj^ transmitted through heredity to the descendants of a given ancestral pair are acquired during the lifetime of these indi- viduals, whereas in the latter the transforming increment is a merely accidental modification arising from unknown causes and hence called spontaneous. The theorj^ is that such .spon- taneous variations are constantly taking place in all individ- 22 BIOLOaiCAL SOCIETY OF WASHINGTON. uals, some in one direction and some in another, and that all except the advantageous ones are immediately lost, while such as tend to increase the chances of survival in the strug- gle for existence are preserved. Nature has thus provided, through this survival of the fittest, for the maintenance of the equilibrium between the organism and the environment, and also for the increase of structural adaptation and vital power, independently both of the effort of the individual to conform more exactly to its surroundings and of the reaction of the organism upon the impinging environment. There has never been anj- doubt of the perfect transmissi- bility of these spontaneous modifications, or, as they have been called, fortuitous variations. They belong to the essential na- ture of the organism, and have, as we shall see later on, been ingeniously explained as originating in the very germ itself. But with regard to functional modifications, or as they are more commonly called, acquired characters, grave doubts have arisen in the minds of many natiiralists as to whether they are capable of being inherited by the descendants of those in which they have been superinduced. They are in a certain sense for- eign to the organism, external and superficial, and the great question has been how they can succeed in so affecting the reproductive germs of the parents as to reappear in the off- spring. That Darwin believed in the transmission of func- tionally acquired characters is attested not only by many passages in which this belief is expressly stated but by the bringing together by him of more facts in support of it than have been given by all other writers combined either before or since. And although the greater part of his work was naturally directed to the establishment of the hitherto un- known, but as he believed, more important law of selection, nevertheless Darwinism proper must be made broad enough PRESIDENTIAL ADDRESS. 23 to embrace both of the great agencies of organic transforma- tion, the functional and the selective. It is hardl}^ necessary to add that pure Lamarckism has nothing whatever to do with such a question as whether acci- dental modifications produced upon an organism, such as mu- tilations from whatever cause, are inheritable, since these are not due to continuous interaction between organism and envi- ronment, are not the objects of the creature's efforts, and are not acquired by any functional or habitual activities. And yet it is no exaggeration to say that at least one-half, probabl}' much more, of the space devoted by the Neo-Darwinians to the sup- posed refutation of Lamarckism has been directed to proving that acccidental mutilations are not transmitted to offspring. I do not deny that there is a doctrine of the transmissi- bility of mutilations, and Darwin and others have collected a large body of facts pointing strongly in that direction, while Brown-Sequard is believed by many to have demonstrated that hereditary epilepsy may be artificially superinduced in guinea-pigs by lesions of the brain. And it may be that Lamarck, coming upon similar facts, gave them a certain cre- dence, but, as we have seen from typical passages quoted from his work, these cases are not capable of being used in support of his general philosophy, which rests entirely upon the effects of functional activities exerted in response to secular alterations in the surrounding conditions of existence. Whatever of truth, therefore, there may be in the doctrine of the transmissibility of suddenl}^ or accidentally acquired characters, it is clearly outside the present discussion and need not be further touched upon. After the doctrine of natural selection had been clearly ex- plained it was found to be so simple and at the same time so far-reaching that it began to be questioned whether much that 24 BIOLOGICAL SOCIETY OF WASHINGTON. had been formerly attributed to the other agency ought not to be credited to it instead ; and it cannot be denied that this in- quiry tended to broaden the field of the selective at the ex- pense of that of the functional principle. So clear and certain are the workings of the former that it is considered safe to credit it with every fact that can be explained by it, even though it be also explicable by the other law. But it was not allowed to rest here. The difficulties in the way of accounting for the transmission of qualities originating after the birth of the parents appeared to some so great that they began to doubt whether in fact such a thing is really possible. Of course, there were many popular and superficial writers on evolution who fail to distinguish the two principles and talked as though all development was due to natural selection, so that to the unscientific and popular mind evo- lution and natural selection were largely synonymous and vaguely comprehended, as is, in fact, still to a large extent, the case. Other better informed people, including some naturalists of note, were so dazzled by the new idea that they lost sight of the old one, and habitually ignored the func- tional element without criticising it or taking any account of it. It appears to have been against this class that Mr. Her- bert Spencer's brilliant exposition of the principle which, in characteristic language, he calls "direct equilibration" was directed. To this I shall have occasion to revert. For the present I propose to confine myself to those writers who clearly comprehend the nature of the two principles, and who either gravely doubt for what seem to them sufficient rea- sons, or else deny altogether the efficacy of functional modifi- cation and the doctrine of the transmission of acquired charac- ters. The limits of an address such as this preclude any effort to make the discussion historically exhaustive by enumerating PRESIDENTIAL ADDRESS. 25 all the investigators who from the first to last have taken this view, or some modified form of it, and I shall be content to name among Germans Du Boise Reymond, Pfliiger, His, and Weismann, and among Englishmen Galton, Wallace, and Ray Lankester ; while what I shall have time to say relative to the nature of the objections raised by these authors will be chiefly confined, for the present, to the views of Galton and Weismann. THEORIES OF HEREDITY. It must, however, be premised that inasmuch as the objec- tions raised against the doctrine of the transmission of ac- quired characters are based upon the difficulties encountered in attempting to explain how such characters can impress themselves upon the germ, all those who have doubted or de- nied such transmission have approached the subject from the side of embryology, which makes their arguments difficult to explain to biologists in general and still more so to the general public. The laws and processes of heredity are still in the stage of mystery, and their mysterious character has led to many erroneous beliefs and popular superstitions. It is a sig- nificant fact that all the mysteries that have been thus far cleared up by science — astronomical, physical and chemical mysteries — have been shown to be the expressions of previ- ously unknown laws of matter and force, and to rest upon a purely material and mechanical basis. The chief obstacle to their comprehension has been the minuteness of the material elements in action — a minuteness far beyond the capacity of the most powerful artificial aids to the senses — so that their secrets have had to be wrung from them by ingenious and multiplied experiments upon their effects. Now, the ultimate 26 BIOLOGICAL SOCIETY OF WASHINGTON. reproductive elements, though doubtless many times larger than any chemical molecule, even the most complex, such as those of protein and other organic compounds, are doubtless still far too minute to be observed by the highest powers of the microscope, and if the entire history of the formation of a new organic being is ever to be learned it must be by a suc- cessful study of the actions of such minute objects. But this is infinitely more difficult than the study of the actions of inorganic elements, since they take place within an organism whose destruction destroys their vital character. In view of the history of the less complex sciences it is natural that biologists should insist that the phenomena of heredity are due to the activities of the ultimate material reproductive elements, and not to any vague and occult force or deus ex niachina. Consequently we find that the only theories of heredity that have been put forth have been based on this assumption. One of the earliest, and certainly the most celebrated of such theories is Darwin's pangenesis, published in 1867.* Accord- ing to this theory, which is doubtless familiar to most of you, the ultimate reproductive elements, called gem mu/es, are given off from the cells of all parts of the body and collect in the germ-cells and sperm-cells, so that the fertilized ova contain literal representatives of every organ and every part of both parents, which in the new being return to their respective locations and cause the repetition in each of the exact qualities possessed by the parental organs or parts, subject, of course, to the modifications due to a conflict or cooperation between the genimules of the two parents, equalizing a character where they are different, and emphasizing it where they are alike. * Variations of Animals and Plants under Domestication. Vol. II, Chaps. XXXVII, XXXVIII. PRESIDENTIAL ADDRESS. 27 It will be readily seen that this theory adapts itself to the broadest conception of heredity and, if trne, accounts for the transmission of functionally produced modifications as well as the selection of such accidental ones as prove advantageous. But to the ordinary mind this strictly materialistic explanation of heredity seems crude and is to a large extent unintelligible, and the doctrine of pangenesis has gained few adherents among scientific men. They fail as a rule to comprehend Mr. Darwin's gemmules and to understand how they should behave in the manner required by the theory. \^ery much of this difficulty, however, is cleared away by the admirable exposition of Mr. Herbert Spencer of the na- ture of what he calls physiological units. To the biologist the organic unit is the cell and when he has explained the nature and action of cells he thinks he has gone far enough. But the facts of heredity cannot be explained by any phenomena man- ifested b}^ cells. Between the cell or morphological unit in biology and the molecule of a highly complex organic com- pound, such as albumen, — the highest class of chemical units — no intermediate element had hitherto been recognized. Mr. Darwin's gemmule is clearly such an intermediate element, and the question at once arose, is there any such ? Mr. Spen- cer has, I think, shown beyond the possibility of doubt that there is and must necessarily be such an element — a unit which is not chemical, since it possesses life, and which is not the morphological unit or cell, bvit is that of which the active part of every cell consists, and is appropriately termed the physio- logical unit. I have elsewhere* undertaken to show that life may have resulted from a process of chemical recompounding, *Americau Naturalist, Vol. XVI, December, 1882, p. 976. Dynamic Sociology, New York, 1883, Vol. I, p. 311. 28 BIOLOGICAL SOCIETY OF WASHINGTON. and may actually constitute the leading property of the high- est organic compound protoplasm, and I venture to suggest here that the gemmules of Darwin and the physiological units of Spencer may be nothing more than the molecules of protoplasm, which, as I have explained, are so immensely complex that any required degree of difference in their essen- tial constitution may easily exist. The only other theory of heredity which time will warrant my mentioning now is that of Professor Haeckel, published in 1876 and known as "the perigenesis of the plastidule." To avoid the possibility of misstatement, I will give this theory in the words of its author, as epitomized in the latest (8th) edition (1889) of his Schopfungsgeschichte (pp. 200-201): "The perigenesis-theory was founded by me in 1876 in a memoir ' on the wave-reproduction of vital particles or the perigenesis of plastidules,' and as a ' provisional attempt at a mechanical explanation of the elementary processes of devel- opment,' and especially of heredity. (In the second part of my collection of popular lectures, Bonn, 1879, pp. 25-So). The perigenesis-hypothesis seeks to explain heredity by a simple mechanical principle, namely, by the well-known principle of transmitted motion. I assume that in everj^ process of repro- duction not only is the special chemical composition of the plasson or plasma transmitted from the parent to the offspring, but also the special form of molecular motion which belongs to its physico-chemical nature. In harmony with the funda- mental laws of modern histology and histogeny, I assinne that this plasma (either the caryo-plasma of the cell-nucleus or the cytoplasma of the cell-body) is alone the original bearer of all vital activity, and hence also of heredity and reproduction. In all plastids (as well the anucleated cytodes as the genuine nucleated cells) this plasma or plasson is composed of plasti- PRESIDENTIAL ADDRESS. 29 dules or plasma-molecules, and these are ' probably sur- rounded by aqueous envelops ; the greater or less thickness of these aqueous envelops, which at once separate and bind the neighboring plastidules, determines the softer or harder condition of the flowing plassou' (p. 48). ' Heredity is the transmission of plastidule motion, whereas adaptation is change of plastidule motion' (p. 55). This motion may in its general aspects be conceived as a ramified wave-motion. In all protists or unicellular organisms (proto- phytes and protozoans) this periodical movement of the mass goes on in a correspondingl}' simple manner while in all tissue-bearing or multicellular creatures (metaphytes and metazoans) it is combined with a mutual generation of the plastids and a division of labor of the plastidules." It will be observed that although this theory of heredity lays special stress upon the idea of motion, thereby recogniz- ing the element of force, it is nevertheless based like all others upon the existence of ultimate material elements different on the one hand from the chemical molecules and on the other from cells, and intermediate between these. The gemmule of Darwin, the physiological unit of Spencer, and the plastidule of Haeckel are the same in essence, and the study of the phe- nomena of these ultimate elements of biology open up a new and most promising field of research into which scarcely any investigator has as yet deliberately entered. We are now prepared to consider the objections of Galton and Weismann to the doctrine of the transmission of function- ally acquired characters. VIEWS OF MR. GAtTON. The earliest expression of Mr. Galton's views, so far as I am aware, is contained in a paper "On Blood-Relationship " 30 BIOLOGICAL SOCIETY OF WASHINGTON. presented by him on June 13, 1872, to the Royal Society of lyOndon and published in its proceedings.* In this paper stress is laid upon the distinction in embryonic development between what he calls the "patent" and the " latent " ele- ments, and he argues from the facts of reversion and atavism that the greater part of the parental elements in heredity are latent in the germ, but prepared to express themselves in more or less remote decendants. Although he addres.ses himself to the anthropologist rather than the biologist, and claims only to be making a contribution to the difficult sub- ject of kinship, he nevertheless evinces a clear grasp of the embryonic conditions of the problem, and as we shall see, anticipates, some of the more exact conceptions of Weismann. He does not wholly deny the possibility of the transmission of acquired characters, but says that " the effects of use and disuse of limbs, and tlio.se of habit, are transmitted to pos- terity in only a very slight degree." In this respect Mr. Galton makes only a slight advance toward the conclusions of Weismann in the much more elabo- rate paper which he read before the Anthropological Institute of Great Britain on November 9, 1S75, and which appeared in the December number or the Contemporary Review for that year, and also in an expanded form in the Journal of the Insti- tute (Vol. V, p. 329). In this paper which is entitled "A Theory of Heredit3%" he, however, approaches the main ques- tion with much greater directness. "The facts" he says "for which a complete theory of heredity must account' may con- veniently be divided into two groups ; the one refers to those inborn or congenital peculiarities that were also congenital in one or more ancestors, the other to those that were not con- genital in the ancestors, but were acquired for the first time by *Vol. XX, p. 394. PRESIDENTIAL ADDRESS. 31 one or more of them during their lifetime, owing to some change in the condition of their life. The first of these two groups is of predominant importance, in respect to the number of well- ascertained facts that it contains, many of which it is possible to explain in a broad and general way, by more than one theory based on the hypothesis of organic units. The second group includes much of which the evidence is questionable or difficult of verification and which, as I shall endeavor to show, does not for the most part, justify- the conclusion commonly derived from it." He further says that his theory " is largely based on the arguments and conditions brought forward by Mr. Darwin in support of pangenesis ; nevertheless the conclusions in this paper w^ill be seen to differ essentially from his own. Pan- genesis appears more especially framed to account for the cases which fall in the second of the above-mentioned groups which are of a less striking and assured character than tho.se of the first group, and it will be seen that I accept the theory of pan- genesis with considerable modification, as a supplementary and subordinate part of a complete theory of heredity, but by no means for the primary and more important part." • He employs the term s//rp " in a special sense — to express the sum-total of the germs, gemmules, or whatever they niaj^ be called, which are to be found, according to every theory of organic units, in the newly fertilized ovum — that is in its earliest pre-embryonic stage." In defending the theory of organic units he says : "We must bear in mind that the alternative hypothesis of a general plastic force resembles that of other m3'stic conceptions current in the early stages of many branches of physical science, all of which yielded to molecular views, as knowledge increased." The paper is an exceedingly luminous contribution to the subject, and the theory advanced may be designated in general 32 BIOLOGICAL SOCIETY OF WASHINGTON. terms as the doctrine of natural selection or survival of the fittest among the organic units constituting the stirp, to de- termine which shall become manifest in the offspring and which shall lie latent to reappear or not in later generations. As the stirp contains organic units that have lain latent in previous generations and may become patent in the generation in ques- tion, the theor}- accounts for reversion, atavism, and the whole train of facts in heredity that have so long puzzled the scientific investigator. We are at present onl}- concerned with so much of it as relates to the transmission of acquired characters. The following passage expresses his views on this point : " We have thus far dealt with three agents — (i) the stirp, which is an or- ganized aggregate of a host of germs ; (2) the personal struc- ture, developed out of a small portion of these germs ; and (3) the sexual elements, generated by the residuum of the stirp. The cases before us are those which are supposed to prove that 2 reacts on 3 — that is, the personal structijre upon the .sexual elements. The first and largest class of these cases refer to adaptivity of race. It is said that the structure of an animal changes when he is placed under changed conditions ; that his offspring inherit some of his change ; and that they vary still further on their own account, in the same direction, and so on through successive generations, until a notable change in the congenital characteristics of the race has been effected. Hence it is<:oncluded that a change in the personal structure has re- acted on the .sexual elements. For my part, I object to so general a conclusion." And he proceeds to elaborate his reasons for such objection. Passing over these for want of time I will conclude this exposition of Galton's views by quot- ing the following passage : "The conclu.sion to be drawn from the foregoing arguments is, that we might almost reserve our belief that the structural PKESIDENTIAI. ADDRESS. 33 cells can react on the sexual elements at all, and we may be confident that at the most they do so in a very faint degree ; in other words that acquired modifications are barely, if at all, inherited, in the correct sense of that word. If they were not heritable, then the second group of cases would vanish, and we should be absolved from all further trouble about them ; but if they exist, in however faint a degree, a complete theory of heredity must account for them. I propose, as alread}^ stated, to accept the supposition of their being faintly heritable, and to account for them by a modification of pangenesis." I am not aware that l\Ir. Galton has modified the views here expressed since the date of that paper, but in all his subsequent ones, as well as in his work on "Hereditary Genius" (iSyg) he continues to emphasize the paramount importance of the latent elements in heredity, and the superiority, as he forcibly expresses it, of nature over nurture. TEACHINGS OF PROFESSOR WEISMANN. The vigorous onslaught which has been made upon the doc- trine of the transmission of acquired characters, since the date of Mr. Galton 's papers, and apparently without a knowledge of them, by Prof. August Weismann of the University of Frei- burg has probably aroused a greater amount of interest among scientific men than any other event that has transpired since the appearance of Darwin's Origin of Species. Professor Weis- mann is an embrjologist and histologist and has conducted a series of prolonged and successful investigations upon several groups of lowly organisms. But he has looked beyond the special facts which are immediately connected with his re- searches and has thought out for himself all the deeper prob- lems of biology. Besides making himself complete master of 34 BIOLOGICAL SOCIETY OF WASHINGTON. the whole field of that science as generally accepted he has co- ordinated its facts and drawn from them a nnmber of new and brilliant conclusions which have set the world to work on entirely new lines of investigation. Professor Weismanu was logically led to the conclusion that acquired characters cannot in any conceivable way be trans- mitted. The first of the series of essays which have produced such a sensation, that on the duration of life, was originally read before the Association of German Naturalists and Physi- cians at Salzburg in September, 1 88 1, and a short abstract of it appeared in Nature for April 5. 1888 (Vol. XXXVII. p. 541). It was in this paper that he elaborated the theory that unicellular organisms are potentially immortal. The second of the series, that on heredity, was his inaugural agldress as Pro-rector of the University of Freiburg, delivered June 21, 1883. It was in this that he first attacked the doctrine of the transmission of acquired characters, and in it and the preceding essay may be found the germs of all his later theories. The remaining six essays appeared at intervals from 18S3 to 1S88. Abstracts and reviews of them occured in Nature and the English magazines, and long before the appearance in 1889 of the admirable work containing an English translation of the whole series with numerous additions and amendments by the author and notes by the translators, "^ the controversy had be- gun in which so many of the most eminent biologists of Europe and America have taken part. Professor Weismann's general course of reasoning is some- what as follows : It is universally admitted that all the higher organisms consist of tissues made up of cells and that these * Essays upon Heredit}' and kindred biological problems. By August Weismann. Authorized translation edited by Edward B. Poulton, Selmar Schonland, and Arthur E- Shipley. Oxford, Clarendon Press, 1889. PRESIDENTIAL ADDRESS. 35 cells do not differ essentially from those which are found lead- ing an independent existence and are termed unicellular or- ganisms. Many of these unicellular organisms reproduce by the process known as fission or division ; that is, they split or divide into two equal parts each of which becomes a new or- ganism exactl}- like the original. These halves exist for an appointed time, increase in size until they are each equal to the original cell before division, and then divide again, so that what was formerly one now becomes four. Each of these four repeats the process, and so on, thus multiplying in a geometri- cal ratio. But if we follow any one of these lines of descent we observe that the last of the line contains some of the same mat- ter that was in the first, and none of the matter has ceased to live. Unless destroyed by some external cause all of the sub- stance of the original cell will continue to live for any conceiv- able length of time. It is " potentially immortal." Now, the theory of descent as a universal organic principle, which Weis- mann fully accepts, explains all the life of the globe as result- ing from previous life through some form of reproduction. Fis- sion is the simplest form of reproduction, and it is found that it is the common form of cell-reproduction within the tissues of the higher animals. All growth is brought about by it or some modification of it. A study of the phenomena of reproduction in the lower organisms shows that it takes place ultimately through some similar process, which, however greatly modi- fied in its details, consists essentially in the actual transmission of the reproductive cell-substance from parent to offspring, and Weismann maintains that the reproductive cells, like those of unicellular organisms, are immortal or perpetual, and that nothing can get into the body of the offspring except through that of one or other of its parents. This is his fundamental doctrine of the continuity of the germ-plasm. The impregnated 36 BIOLOGICAL SOCIETY OF WASHINGTON. ovum contains the germ-plasms of the two parents, and out of it the embryo is formed. The embryo develops independently of the mother by a circulation of its own, and no external in- fluences can b}' any conceivable method affect or change the characters of the offspring. But it is well known that variation takes place, that the off- spring does not always resemble either parent, and that changes go on so great as to result in the creation of new species, new genera, and entirely new types of life. All this Weismann ad- mits. How does he explain it ? Primarily by natural selec- tion, but he does not stop with that. It has always been ad- mitted that natural selection did not explain the cause of varia- tion. Weismann attempts to do this, and his reasoning is exceedingly ingenious. The original reproductive cells are assumed by him to con- sist of an indefinite number of units which he calls germ-plasms, and their presence is explained on the assumption of their pre- servation from ancestral organisms. Asexual reproduction is of course incapable of producing variation, and he maintains that sexual reproduction has been developed and exists solely for the purpose of insuring variation. Relative to the constitution of the germ-plasm he says : " Every detail in the whole organism must be represented in the germ-plasm by its own special and peculiar arrangement of the groups of molecules (micellae of Nageli) and the germ-plasm not only contains the whole of the quantitative and qualitative characters of the species, but also all individual variations as far as these are hereditary : for example the small depression in the center of the chin noticed in some families. The physi- cal causes of all apparently unimportant hereditary habits or structures, of hereditary talents, and other mental peculiarities, must all be contained in the minute quantity of germ-plasm PRESIDENTIAL ADDRESS. 37 which is possessed by the nucleus of a germ-cell ; not indeed, as the preformed germs of structure (the gemmules of pangene- sis), but as variations in its molecular constitution ; if this be impossible, such characters could not be inherited " (pp. loo, lOl). The union of two germ-cells from entirel)^ different indi- viduals always multiplies the number of ancestral germ-plasms by two. The excess is kept down by the removal of the second polar body, as he supposed was proved by its not taking place in parthenogenesis. But the part removed as w^ell as the part retained contains germ-plasms from both parents alike and hence the offspring must partake of the nature of both. These ancestral germ-plasms exist in the reprod'uctive cells in vast numbers, and in the removal of half of them at each union of the sexes, there must remain not merely those of the immediate parents, but those of previous generations. If we were theoretically to conceive that at the outset only a single germ-plasm existed from each parent, then the second genera- tion would transmit four, the third eight, and so on in a geometrical ratio, until they would become so numerous as to require the removal of a portion and ultimately always of half the ancestral germ-plasms at each act of reproduction. Says Professor Weismann : " These different qualities are what I have called the ancestral germ-plasms, i. e., the germ- plasms of the different ancestors, which must be contained in vast numbers, but in a very minute quantities, in the nuclear thread. The supposition of a vast number is not only required by the phenomena of heredity but also results from the com- paratively great length of the nuclear thread ; furthermore it implies that each of them is present in very small quantity. The vast number together with the minute quantity of the ancestral germ-plasms permit us to conclude that they are. 38 BIOLOGICAL SOCIETY OF WASHINGTON. upon the whole, arranged in a linear manner in the thin thread- like loops ; in fact the longitudinal splitting of these loops appears to me to be almost a proof of the existence of such an arrangement, for without this supposition the process would cease to have any meaning" (pp. 359-360). His general view of the origin of variation is thus given by him: "It is well known that this process [sexual or amphi- gonic reproduction] consists in the coalescence of two distinct germ-cells, or perhaps onh' of their nuclei. These germ-cells contain the germ-substance, the germ-plasm, and this again, owing to its specific molecular structure, is the bearer of the hereditary tendencies of the organism from which the germ- cell has been derived. Thus in amphigonic reproduction two groups of hereditarj- tendencies are as it were combined. I regard this combination as the cause of hereditary individual (Characters, and I believe that the production of such characters is the true significance of amphigonic reproduction. The object of this process is to create those individual differences which form the material out of which natural selection produces new species " (p. 272). " I do not know what meaning can be attributed to sexual reproduction other than the creation of hereditary individual characters to form the material upon which natural selection may work" (p. 281). "The most important dut}^ of sexual reproduction is to preserve and continually call forth individual variability, the foundation upon which the transformation of species is built" (p. 373). " Sexual reproduction is to be explained as an arrangement which ensures an ever-varying supply of individual differ- ences" (p. 384). Weismann's classification of cells into somatic and reproduc- tive is fundamental to his whole philosophy. On this point he says : ' ' The first multicellular organism was probably a clus- PRESIDENTIAL ADDRESS. 39 ter of similar cells, but these units soon lost their original homogeneity. As the result of mere relative position, some of the cells were especially fitted to provide for the nutrition of the colony, while others undertook the work of reproduction. Hence the single group would come to be divided into two groups of cells, which may be called somatic and reproductive — the cells of the body as opposed to those which are concerned with reproduction (p. 27) , . . As the complexit}- of the metazoan body increased, the two groups of cells became more sharply separated from each other. Very soon the somatic cells surpassed the reproductive in number, and during the in- crease they became more and more broken up by the principle of the division of labor into sharply separated systems of tis- sues. As these changes took place the powxr of reproducing large parts of the organism was lost, while the power of repro- ducing the whole individual became concentrated in the repro- ductive cells alone " (p. 28). His theory further assumes that the germ-cells contain two kinds of plasm, which he calls re- spectiveh' the ovogenetic and the somatogenic, i. e., the first capable onh- of producing germ-cells, the latter capable only of producing somatic cells. These exist together in the fertil- ized ovum, and if allowed to remain there would go on repro- ducing themselves in something like equal numbers. But the bod}^ consisting almost entirely of somatic cells, it is evident that such a multiplication of germ-cells would be onh- a hin- drance to development. This, he claims, explains the myste- rious phenomena so long observed by embryologists and called the removal of polar bodies. The polar body first removed is nothing more nor less than the ovogenetic nucleo-plasm, which is now in the way, and whose removal is necessary to the formation of the embryo. This is the work alone of the somatic cells, and these, consisting as they do of the germ-plasms of an 40 IJIOLOUICAL SOCIETY OF WASHINGTON. indefinite series of ancestors, and containing representatives of every part of the parent organism, proceed to reproduce a new creature on the hereditary type of the parents with the modifi- cations due to the commingling of many ancestral types. Without dwelling longer upon these ultimate processes which constitute the premises of Weismann's argument, I will now proceed to state his conclusion. It is simply that he is utterl}^ unable to see how the somatic cells of an adult indi- vidual can react upon or in any waj^ affect its reproductive cells. If it cannot, the transmission through either parent to its offspring of any peculiarity acquired since the embryo of the parent began to form is impossible. Firmly believing in the truth of his theory he stoutly insists that no such thing can take place. Of course it needs to be clearly understood what he means by acquired characters, and here, it is claimed lies the chief point in dispute between the Neo-Darwinians and the Neo-Lamarckians. The former contend that the latter class as acquired characters those which are simply due to natural selection. It will therefore be profitable to dwell a moment upon this point. " The tendencies of heredity ", says Weismann, " of which the germ-plasm is the bearer, depend upon this very molecular structure, and hence only those characters can be transmitted through successive generations which have been previously in- herited, viz. , those characters which were potentially contained in the structure of the germ-plasm. It also follows that those other characters which have been acquired by the influence of special external conditions, during the lifetime of the parent, cannot be transmitted at all " (p. 267). " It is only by suppos- ing that these changes arose from molecular alterations in the reproductive cell that we can understand how the reproductive cells of the next generation can originate the same changes in RESIDENTIAL ADDRESS. 41 the cells which are developed from them ; and it is impossi- ble to imagine any way in which the transmission of changes produced by the direct action of external forces upon the somatic cells, can be brought about . . . To this class of phenomena of course belong those acts of will which call forth the func- tional activity of certain groups of cells " (p. 80). " Only those new characters can be called ' acquired ' which owe their origin to external influences, and the term ' acquired ' must be denied to those which whollj^ depend upon the mysterious relationship between the different hereditary tendencies which meet in the fertilized ovum. These latter are not ' acquired ' but inherited, although the ancestors did not possess them as such, but only, as it were, the elements of which they are composed " (p. 252). "If acquired characters are brought forward in connexion with the question of the transformation of species, the term ' acquired ' must only be applied to those characters which do not arise from within the organism, but which arise as the re- action of the organism under some external stimulus, most commonly as the consequence of the increased or diminished use of an organ or part " (p. 322). That such characters cannot be inherited he asserts with the strongest emphasis and frequent iteration. His treatment of this point often borders on the dogmatic, as a few extracts will show. " It has never been proved " he says, " that acquired char- acters are transmitted, and it has never been demonstrated, that, without the aid of such transmission, the evolution of the organic world becomes unintelligible. The inheritance of ac- quired characters has never been proved, either by means of direct observation or by experiment " (p. 81). "No single fact is known that really proves that acquired characters can be transmitted" (p. 267). " If acquired characters cannot be 42 BIOLOGICAL SOCIETY OF WASHINGTON. transmitted the Lamarckian theory completely collapses, and we must entirely abandon the principle on which alone Lamarck sought to explain the transformation of species, — a principle of which the application has been greatly restricted by Darwin in the discovery of natural selection, but which was still to a large extent retained by him. Even the apparently powerful factors in transformation — the use and disuse of organs, the re- sults of practice or neglect — cannot now be regarded as pos- sessing any direct transforming influence upon a species, and the same is true of all the other direct influences, such as nutri- tion,light, moisture, and that combination of different influences which we call climate. All these, with use and disuse, may perhaps produce great effects upon the body (soma) of the in- dividual, but cannot produce any effect in the transformation of the species, simply because they can never reach the germ- cells from which the succeeding generation arises" (pp. 387- 388). And much more in the same strain. Weismann fully admits the influence of the environment upon the individual in producing marked changes. He also fully admits the facts of adaptation to environment and the transformation of species and development of organic beings. But he insists that natural selection is competent to explain all this, that it takes place through the selection of such accidental variations in the germ as prove advantageous, or, as he puts it, the selection from among an infinite number of ancestral germ- plasms in the fertilized ovum of such as will produce an indi- vidual most in harmony with its environment, leaving all others in the latent state. This, as we have seen, is pure Galtonism. But this incapacity for the inheritance of acquired characters is confined to metazoans or multicellular organisms — organisms whose reproductive and somatic cells are differentiated. It does not apply to protozoans or unicellular organisms. These PKESIDKNTIAL ADDRESS. 43 are greatly influenced by the environment, and, consisting en- tirely as it were of reproductive cells, naturally transmit their variations to their descendants directly. Only thus can vari- abilit}^ be perpetuated, and whatever is true of them is true of all germ-cells. "The origin of hereditary individual vari- ability," says Weismann, "cannot indeed be found in the higher organisms — the Metazoa and Metaphj-ta ; but is to be sought for in the lowest — the unicellular organisms. In these latter the distinction between body-cell and gerra-cell does not exist. Such organisms are reproduced b}- division, and if, therefore, any one of them becomes changed in the course of its life by some external influence, and thus receives an indi- vidual character, the method of reproduction ensures that the acquired peculiarity will be transmitted to its descendants ' ' (pp. 277-278). It is here that comes in his fundamental doctrine of the con- tinuit}' of the germ-plasm. If not the germ-cells, at least the germ-plasm of either parent passes intact to the offspring. It is perpetual, or as he calls it, immortal. It gives to the new being its special character, but receives nothing from it. It remains in the offspring until it in turn becomes a parent, and again passes to the third generation without ever having ceased to live. Ever}' living being on the globe to-day contains in its germ-plasm something that has never ceased to live since the original life-breath was breathed into organic nature. Through all the ancestral t3'pes of the phyletic chain it has persisted, passing from parent to offspring through the trans- forming series, so that in the loins of the highest types of man there is something which was still living in the lowest primor- dial worm and even in the bathybian ooze of those primeval waters which in the earliest Cambrian times succeeded the formation of the original crust of the globe. 44 BIOLOGICAL SOCIETY OF WASHINGTON. Upon this series of brilliant speculations and startling asser- tions, including much that it has been impossible for me to bring forward, has been founded the school of Neo-Darwinism. In Germany they attracted comparatively little attention, in France none, but in England they have become almost a shib- boleth in the mouths of a large class of leading biologists. It unfortunately requires something more than mere truth to arouse enthusiasm in many minds, and however much it may be di.sclaimed, it cannot probably be justly denied that the peculiar position of prominence and honor which this theory gives to the doctrine of natural selection, conceived and elab- orated by Englishmen, had much to do with its especial charm for English ears. It is not to be supposed that Weismann de- liberately bid for applause from England, but he could clearly see the tendency of his doctrines to exalt natural selection. He does not allude to this in any of his earlier essays, nor until he had begun to observe the effect his writings were pro- ducing in England. In the preface to liis nfth essay, dated Nov. 22, 1885, however, he says : " The transmission or non- transmission of acquired characters must be of the highest im- portance for a theory of heredity, and therefore for the true appreciation of the causes which lead to the transformation of species. Any one who believes, as I do, that acquired charac- ters are not transmitted, will be compelled to assume that the process of natural selection has had a far larger share in the transformation of species than has been as yet accorded to it ; for if such characters are not transmitted the modifying in- fluence of external circumstances in many cases remains re- stricted to the individual, and cannot have any part in produc- ing transformation" (pp. 252-253). And in the last essay of this series, originally delivered in September, 1888, he further remarks : ' ' But if the transmission of acquired characters is PRESIDENTIAL ADDRESS. 45 truly impossible our theory of evolution must undergo material changes. We must completely abandon the Lamarckian prin- ciple, while the principle of Darwin and Wallace, viz. , natural selection, will gain an immensely increased importance " (p. 423)- A CRITIQUE OF WEISMANN. I have now, as I believe, fairly if not fully stated, chiefly in the language of its founder, the Neo-Darwinian theory, and before passing to consider what has been said on the other side, and the position of the Neo-L,amarckians in general, I would like to pause a moment in order to offer a few reflections of my own upon Weismann's teachings. I am emboldened to do this the more not only because I have not seen the exact point of view from which they especially strike me touched upon by others in the voluminous discussion which has grown out of them, but also because what I shall say will be based entirely upon his own statement of the facts, and therefore the objec- tion that, not being an embryologist, I am not competent to weigh the considerations from that side (which I would freely admit), cannot properly be raised. The question is whether, accepting the continuit}- of the germ-plasm, accepting the nature which he ascribes to the fer- tilized ovum with its multitudes of ancestral plasms out of which selections are made, accepting his explanation of the meaning of the first and second polar bodies, accepting his differentiation into reproductive and somatic cells, and all the other details which he brings forward, many of which are, of course, only hypotheses, there do not still remain grounds on which to base a theorj^ of the transmission of certain kinds of acquired characters, and especially those of a strictly functional nature. In fact, the question seems to me rather to be whether 46 BIOLOGICAL SOCIETY OF WASHINGTON. his line of argument carried to its extreme logical conclusion would not preclude the possibility of any variation whatever even in the germ-plasms themselves. It is not sufficient to say that all variation is due to the varied character of multitudin- ous germ-plasms in the fertilized ovum, brought there from many often remote ancestors possessing ver}- different charac- ters. This is a pctitio principii, since it assumes these differ- ences in those ancestors, and the primary question must be answered ; whence these ancestral differences ? How does he account for any differences at all ? We have already seen that Weismann restricts his denial to multicellular organisms and admits as a necessary part of his theory, that unicellular organisms are easily affected by the nature of their surroundings and activities, and that the changes thus produced are directly transmitted. "If for in- stance," he says, "a protozoan, by constantly struggling against the mechanical influence of currents in water, were to gain a somewhat denser and more resistent protoplasm, or were to acquire the power of adhering more strongly than the other individuals of its species, the peculiarity in question would be directly continued on into its two descendants, for the latter are at first nothing more than the two halves of the former. It therefore follows that every modification which appears in the course of its life, every individual character, however it may have arisen, must necessarily be directly transmitted to the two offspring of a unicellular organism (p. 278). . . . We are thus driven to the conclusion that the ultimate origin of hereditar}^ individual differences lies in the direct action of ex- ternal influences upon the organism " (p. 279). But he even goes further and asserts that there is no other waj^ b}' which the germ can be affected. " I have never doubted" he says, ' ' about the transmission of changes which depend upon an PRESIDENTIAL ADDRESS. 47 alteration in the gerni-plasni of the reproductive cells, for I have ahA'a5^s asserted that these changes, and these alone must be transmitted (p. 410) . . . In what other way could the trans- formation of species be produced, if changes in the germ-plasm cannot be transmitted ? And how could the germ-plasm be changed except by th& operation of external influences, using the words in their widest sense?" (p. 411). Now if, as he insists, external influences cannot possibh' af- fect the germs of metazoans, and if, as he here maintains, it is external influence alone that can influence any germs, it must follow that the onh^ variation that could have taken place in the germ-plasms of the highest animals are those which oc- curred in the protozoan stage of their development. This is clearl}- a reductio ad absiirdiwi^ derived entirelj^ from his own statements, some of them among his latest utterances. The difficulty is to see why he should adhere so tenaciously to the idea that the germ-cells cannot be influenced by functional changes in the organism containing them. The mere fact that they are lodged within the body of an animal does not affect the question unless it can be shown that they are so lodged that no change is possible in the nature of their immediate surround- ings. To assume this is gratuitous and contrary to what would be naturally supposed. In reading certain passages in his own book one is strongly tempted to doubt whether he believes it himself. For example, he says in one place speaking of hered- itary variations : "I believe however that thej^ can be referred to the various external influences to which the germ is exposed before the commencement of embryonic development. Hence we ma}' fairly attribute to the adult organism influences which determine the phjdetic development of its descendants. For the germ-cells are contained in the organism, and the external influences which affect them are intimatel}^ connected with the 48 BIOLOGICAL SOCIETY OF WASHINGTON. state of the organism in which they lie hid. . . It is even pos- sible that the effects of these influences may be more special- ized ; that is to say, that they may act only upon certain parts of the germ-cells " (pp. 103-104). But he seems to see a great difference between this and the transmission of characters ac- quired in certain special organs to the sa'me organs of the off- spring. This would probably be clear only to an embryologist. One of the most suggestive thoughts in his whole philosophy is that of the total dissimilarit}^ between the germ and the devel- oped organism which is to result from it. He maintains with every semblance of truth that there can be nothing in common l^etween them except the fact that the molecular structure of the germ is such that if allowed to develop it will produce a being similar to the one from which it sprung. This principle seems to be peculiarly applicable to the subtle influences which effect heredity, and without appealing to anything occult or abandoning the strictly casual and mechanical theory of hered- ity, it may be submitted whether we know enough about it as yet to assert that influences affecting the parental organism, even any of its organs, may not react specifically and in kind upon the germ and set up molecular tendencies in the same direction. This may be said quite independently of any at- tempt to explain precisely how it can do so, as the theory of pangenesis claims to do. If the germ-plasms vary within the body of either parent be- fore they are brought together that variation must be due to influences acting upon them in the animal body. All this Weismann admits, but he denies that the changes which he admits to take place in the individual as the result of changes in the environment and subsequent changes in the habits and activities of the creature can be regarded as among the causes which produce changes in the germ-plasm. Is this logical or PRESIDENTIAL ADDRESS. 49 even reasonable ? If not due to such changes to what causes are they due ? Without pretending to explain how such a thing could happen, I claim that the indications are that it does hap- pen. To say without proof that it cannot happen adds nothing to the argument. We have an antecedent and we have a con- sequent. Both are facts. There is no possibility in the present state of our knowledge of either proving or disproving the casual connection between these facts. Variation takes place in the direction of adaption to changed conditions and activities. So far the inference is confirmed by a third fact. If the inference had not been challenged in the interest of another principle this would be regarded as proof. I do not agree with.Weis- mann that the burden of proof rests on those who draw this natural inference. It rests on him and the Neo-Darwinians to show that the assumed cause is not a cause. This they have thus far failed to do. You will understand that I am speaking of variations which take place in the germ-cells and sperm-cells of parental organ- isms before they blend in the fertilized ovum. Most of Weis- mann's argument is directed to show that the fertilized ovum itself cannot be affected by any transforming influence acting upon the mother during the growth of the embryo. This may be true but it is unimportant. The time required to^ develop the embryo is too short for the environment to produce any material change however strongly the tendenc}^ might be at the time in the direction of such change. It is chiefly the un- combined sexual elements which are admitted by all to be un- dergoing specific transformation. The Neo-Darwinians deny that this is due to admittedly parallel transformations going on in the individual, the result of external and internal influences upon the developed body ; the Neo-I^amarckians consider the latter as in great part the cause of the former, while admitting 50 BIOLOGICAL SOCIETY OF WASHINGTON. that Other variations are taking place clue to unknown causes and that these are seized upon by natural selection to the advantage of the species. The difficulty, on Weismann's theory, of accounting for any variation at all above the protozoans still confronts us. If ex- ternal influences can only act on unicellular organisms in such a way as to be transmitted, it must follow that so soon as the multicellular stage is reached a rigid fixity must result. One of these lower metazoans may undergo important modifications during its lifetime, but its offspring are always set back to pre- cisely the same place where the parent was when it set out. All these functionally produced changes are, according to him, utterly lost because they cannot react upon the germ-plasm. Where is the room for the action of natural selection ? He has not dwelt upon this point, but he would probably say, though contrary to statements above quoted, that the germ-plasms are constantly undergoing spontaneous variation and that natural selection works on these. We would then be brought back to where we were a moment ago, with the question still before us, how spontaneous variations differ from functional ones (for he would not maintain that they were wholly uncaused effects), and why it is not logical and rational to assume that functional changes are impressed upon the germ-cells in ways which, though unknown to us, are no more unknown than is the cause of spontaneous variations. This seems to be far more reason- able than the far-fetched, and, as it seems to me, childish view recently expressed by Prof. E. Ray Lankester, that the envi- ronment does indeed influence the germ-cells but only by kaleidoscopically shaking up their contents, thus causing what are called "sports " in the progeny, and that natural selection seizes upon the.se, thereby securing advantageous tran.sfor- mations. TTvESIDENTIAL ADDRESS. 51 NEO-DARWINISM.-^ We will next briefly pass iu review the extraordinar}^ dis- cussion which has followed chiefly from the publication of Weismann's essays. As already remarked, they produced very little influence upon the German mind, and most German in- vestigators who noticed them at all, either saw little in them, or else attacked them with greater or less violence. It is almost exclusively in England that thej^ have found favor, and here a veritable school of biologists has sprung into existence prepared to defend even the most extreme of Weismann's theories. It is due to the German investigator to say that, with the exception of the slight tendency above pointed out to dogmatize on the subject of the non-transmissibility of acquired characters, his essays are dignified and courteous and often evince an almost Darwinian modesty wdth regard to his own theories. Far different was the case with most of his English disciples. What he states as probable they assert as forever settled, and his working hypotheses become for them the funda- mental truths of science. His papers were translated and re- viewed, usually in an aggressive manner before au}^ one had ventured to criticise them. Being usually beyond the reach of an}' but the embryological specialist all except ardent disciples reserved their judgment and declined to enter the field. At first there was an attempt to make it appear that Weismann's views reflected only those of Darwin himself and that all out- side of them consisted in deviations and wanderings from his doctrines. It was sought to stamp them with the name of * The expression Neo-Darwinian was first used, so far as I am aware, by Dr. G. J. Romanes in a letter to Nature for Aug. 30, 1888 (Vol. XXVIII, p. 413), and occurs frequently iu subsequent discussions. The substantive form Neo-Darwinism was a natural outgrowth from it. 52 BIOLOGICAL SOCIETY OF WASHINGTON. "pure Darwinism,"* and the reader was frequently informed what Darwin reall}- intended to say in certain passages which could not otherwise be made to harmonize with the new doc- trine, and even in some still more refractor}^ passages we are told what we would have said "if it had occurred to him." f In default of any real opponent the Duke of Argyll, with his strong theological bias, his medieval spirit of logomachy, and his total lack of scientific ideas, was called out and set up as a sort of man of straw to be repeatedly demolished. But like the shadows in the valley of Walhalla, he emerged each time unscathed and renewed the deathless struggle. His presence in the arena had the further advantage for the new school of affording them an opportunitj^ to point to him as a sample of the opponents of Weismann. Against all this a few protests were raised from time to time and after the appearance of the English edition of the essays a few able and critical analyses were made. But the general character of the discussion as it has gone on in the columns of Nature and in the British magazines is such as I have de- scribed. The only other prominent or frequent contributor in answer to the disciples of Weismann is Dr. G. J. Romanes, and he has been more especially concerned with defending his priority to the idea which he has elaborated under the name of Physiological Selection, and to the discussion of certain phases of the law of panmixia which he claims to have dis- covered. It would, however, be unjust to deny that the dis- cussion has been of value to science, since, had it done no more than to attract wide attention to so momentous a question it could not have been without its uses. '^Nature, Vol. XXXVIII, Aug. i6, 1888, p. 364 ; Aug. 23, 1888, p. 388 ; Vol. XL, pp. 567, 619. t See Nature, VoL XLI, March 27, 1890, pp. 487, 48S. PRESIDENTIAL ADDRESS. > 53 NEO-LAMARCKISM.* Let us inquire what has really been done from first to last toward the demonstration, or scientific establishment of the law of transmission of functionally acquired characters and the preservation through heredity of the modifications produced by changes in the environment. It will not be necessary to go back to Lamarck as his presentation of the subject has been sufficiently dwelt upon. But I cannot agree with some recent writers that Lamarck was defending a totally different prin- ciple from that which is being defended to-day. It is true that Neo-Lamarckians recognize natural selection as an equalh\ and in some respects far more potent law, although, as has been justlj^ insisted upon, it does not explain the cause of the variations of which it makes use. The Lamarckian principle does this, so far as it goes, and affords a true mechanical, itoidea . . . xvii Coccidiiim bigetniimm xviii Color of flowers in attracting insects . . . xiv Color of human hair, change in vi Color of plumage of birds, change in . . . vi Colors of fishes vii Colors of insects vii Columellar plaits, development of, in Gastropods g Committees iji 74 ETOr.OdlCAL SOCIETY OF WASHINGTON. PAGE Cotnmittees for 1S91, announcement of xii-xiii Conscious effort, relations of 3 Cope, K. D 60, 61 C(3;<'//a, a new genus of Compositse . . ix Council iii Co%'ille, F. v.— The new arrangement of genera in the Herbarium of the De- partment of Agriculture v Fruiting of the Ginkgo at the Depart- ment of Agriculture x Food plants of the Indians of the Death Valley region xvii A review of Kuutze's Revisio Generum Plantaruni xix Cretaceous mammals, review of the dis- covery of xiii Crustacean, ostracod, new genus and species of, from Lower Cambrian . . . . v Cuckoo, Asiatic, occurrence of the, on the Pribylov Islands xi Cuckoo stomachs and their contents . xvii Cunningham, J. T 57 Curtice, Cooper. — The moultiugs of the cattle tick (Ixodes bovis) v A preliminary study of ticks in the United States • • . . . . xi Some little known worms in cattle . . . xiii Practical value of investigating parasites of live stock xv Cuvier 19 Cyclopteroidea ix Cypress knees, what are xiv D. Dall, W. H 62 Dall, W. H. — On dynamic influences in evolution vi, i Original drawings of the fur seal and Steller's sea cow vii Paleontological notes from the north- west coast X On the topography of Florida with reference to its bearing on fossil faunas xi Age of the Peace Creek hone beds in Florida xiv Darwin, Charles 12, 13, 18, 20, 22, 23, 26. 31, 45, 53, 61, 65, 66 Darwin, Erasmus 13, 19, 66 Darwinism 12, 19 Date palms, recent introduction of .... xv DeCandolle, Alphonse, on the transmis- sion of acquired characters . ... .xix De Maillet 13 DenWWon oi Desmognathus, notes on . . xvi Desmognathus, notes on dentition of . . . xvi PAGE Dewey, L A., election of xv Dicentta ciicullaria, vegetative propaga- tion of ... X Dichromatism, remarks on xix Direct equilibration . 24,53 Discovery of cretaceous mammals, review of xiii Discovery of vertebrate life in Lower Silurian (Ordovician) strata xiii Discoveries, recent, of Potomac fossil plants near Washington xix Disease, foot, of captive birds x Disease, recent bacteriological progress in prevention and cure of xv Diseases, plant, recent progress in the study of xvi Distribution of animal and vegetable life xiv Distribution of certain mammals, birds and plants on the Pribylov Islands . . . xvi Distribution of fishes by underground water courses xvi Distribution of species, geographical, some early views of viii Drawings, original, of the fur seal and Steller's sea cow vii Du IJois, Raymond 25 Dwarfs, notes on xvi Dynamic evolutionists 2 Dynamic influences in evolution, Dall on vi. i Dynamic variations limited 4,5 E. Echinococcus in swine xviii EchiHorhynrliiis gigas. development of . xvii Edson, J. R., election of viii Effort, conscious, relations of .3 Egleston, N. H.— The temperature of trees xvii Eimer, Theodor 57 Election of Officers xii Embrj'oofchick with two protovertebrae . xiv Entomology, economic, authorship of bibliography of ix Environment .... 54 Environment, its relation to the organism . 2 Environment, selection limited hydjiiain- ics of 6, 9 Equilibration, direct 24, 53 Equilibration, iii