Part 41

The work in systematic zoology is now mainly carried on by specialists in relatively small groups of animals. This is necessitated both by the increasingly large number of species known to science and by the completeness and exactness with which species must now be defined. The majority of systematic workers are now connected with museums where the large collections furnish material for comparative studies.

Prominent in this field is the United States National Museum, the publications of which are mainly taxonomic and zoogeographic, and cover every group of organism. The adequacy of this great museum for such studies may be illustrated by the collection of mammals. This museum has the types of 1135 of the 2138 forms (including species and subspecies) of North American mammals recognized in Miller’s list,[175] and less than 200 forms lack representatives among the 120,000 specimens of mammals. Systematic monographs of several of the orders of mammals have been published.

Systematic study of the birds has brought the number of species and subspecies known to inhabit North and Middle America to above 3000. The most comprehensive systematic treatise is the still incomplete report of Ridgeway[176] of which seven large volumes have already been issued.

On the reptiles, the most complete monograph is that by Cope[177] entitled “The Crocodilians, Lizards and Snakes of North America.”

The Amphibia have also been studied by Cope, whose report on the Batrachia of North America[178] is the standard taxonomic work.

The most comprehensive systematic work on fishes is the “Descriptive Catalogue of the Fishes of North and Middle America” by Jordan and Evermann.[179]

The invertebrate groups have been in part similarly monographed by the members of the U. S. National Museum staff and others, and further studies are in progress. Other taxonomic monographs published by this museum include the various groups of animals from many different parts of the world.

A number of the larger State, municipal, and university museums publish bulletins on special groups represented in their collections as well as articles of general zoological interest.

Expeditions, subsidized by museum and private funds, are from time to time sent to various parts of the world and their results are often published in sumptuous manner.

The total number of living species of animals is unknown, but considering that about a quarter of a million new species have been described during the past thirty years, it is probable that several million species are in existence to-day. More than half a million have been described. These are probably but a small fraction of the number that have existed in past geological ages.

Thus, in spite of all the work that has been done in systematic zoology and as the number of known species continues to increase, there still remain many groups of animals, some of which are by no means rare or minute, in which probably only a small proportion of the species are as yet capable of identification.

It is only since the publication of Ward and Whipple’s “Fresh-water Biology” within the past year that the amateur zoologist could hope to find even the names of all the organisms which may be collected from a single pool of water. And in many cases he will still meet with disappointment, for many of our protozoa and other fresh-water organisms have not yet been described as species.

During the past few years there has been a tendency on the part of some of our biologists engaged in experimental work to disparage the studies of the systematists. It must be granted, however, that both lines of work are essential to the sound development of zoological science, for experimental investigations in which the accurate diagnosis of species is ignored always result in confusion.

_Ecology._—The marvelous modifications in structure and instincts by which the various animals are adapted to their surroundings now forms a special topic in biological research and one of the most fascinating. The adaptations in habitat, time, behavior, appearance and even in structure are found capable of a certain individual modification when studied experimentally.

_Zoogeography._—Closely associated with systematic zoology, and indeed a part of the subject in its broader sense, is the study of the geographical distribution of animal species and larger groups.

_Paleontology._—The geological succession of organisms embraces a field where zoologist and geologist meet. The wonderful progress made by American investigators is well described in the preceding chapters on Historical Geology and Vertebrate Paleontology.

_Biometry._

Since Darwin’s theory of evolution postulated the origin of new species by means of natural selection, it was obviously necessary in order to apply a critical test to determine the precise limits of a species. It was, therefore, proposed to subject a given species to a strict examination by the application of statistical methods to determine the range of variation of its members and the extent to which the species intergrades with others. Other problems, particularly those concerning heredity, were treated in similar manner. This branch of biological science was particularly developed by the English School, led by Sir Francis Galton, followed by Karl Pearson and William Bateson.

In America the methods of biometry have been utilized extensively by Charles B. Davenport, Raymond Pearl, H. S. Jennings and others in the solution of problems in genetics and evolution. Their work shows the great value of critical statistical analysis in the interpretation of biological data. A thorough training in mathematics is now found to be hardly less important for the biologist than is a knowledge of physics and chemistry, for the science of biometry has become one of the most important adjuncts to the study of genetics.

_Comparative Anatomy and Embryology._

_Comparative Anatomy._—Upon the foundations laid down by Cuvier a century ago the present elaborate structure of comparative anatomy of animals, both vertebrate and invertebrate, has been developed. Vast as is the present accumulation of facts and theories many important problems still await their solution. Jeffries Wyman was long a leader in this field, where many workers are now engaged.

_Embryology._—The embryological studies, so brilliantly begun by Von Baer early in the nineteenth century, are still in progress. They have now been extended to the groups more difficult of investigation and into the earliest stages of fertilization and implantation in the mammals. Artificial cultural methods have yielded important results. Louis and Alexander Agassiz, Mark, Minot, Brooks, Whitman, Conklin and E. B. Wilson have taken prominent parts in this work.

In the early nineties embryological studies were directed to the arrangement of cells in the dividing egg, and there was much discussion of “cell lineage” in development. Valuable as were these studies they threw comparatively little light on the general problems of evolution.

_Experimental Embryology._—A more fertile field, developed at the same period and a little later, was found in experimental embryology. The discoveries made by Driesch and others in shaking apart the cells of the dividing egg or by destroying one or more of these cells gave a new insight into the potency of cells for compensatory and regenerative processes. These studies attracted many able investigators, who made still further advance by subjecting the germ cells, developing eggs, embryos, and developing organs to a great variety of artificial conditions.

_Artificial Parthenogenesis._—Another question concerns the nature of the process of fertilization and the agencies which cause the fertilized egg to develop into an embryo. In 1899 Jacques Loeb succeeded in causing development in unfertilized sea-urchin eggs by subjecting them to concentrated sea water for a period and then returning them to their normal environment. To this promising field of experimental work came many of the foremost biologists both in America and Europe. It was soon found that the eggs of most groups of animals except the higher vertebrates could be made to develop into more or less perfect embryos and larval forms by treatment with a great variety of chemical substances, by increased temperature, by mechanical stimuli and by other means. This artificial parthenogenesis, as it is called, has also been successful in plants (_Fucus_), and recently Loeb has reared several frogs to sexual maturity by merely puncturing with a sharp needle the eggs from which they were derived. Loeb, then, maintains that “the egg is the future embryo and animal; and that the spermatozoon, aside from its activating effect, only transmits Mendelian characters to the egg.”[180]

Further experimental analyses of the nature of the fertilization mechanism have recently been made by Morgan, Conklin, F. R. Lillie, and others.

_Germinal Localization._—The question as to whether the egg contains localized organ-forming substances has been studied experimentally particularly by means of the centrifuge. The results indicate that neither of the older opposing theories of “performation” or “epigenesis” is applicable to all eggs, but that in certain organisms the eggs possess a well marked differentiation while in others each part of the egg is essentially, although probably not absolutely, equipotential.

_The Germplasm Cycle._—Since Weismann’s postulation of the independence of soma and germplasm in 1885 many attempts have been made to trace the path of the hereditary substance from one generation to the next. A recent book by Hegner[181] summarizes the success attained in various groups of animals.

_Cytology._

Another important field of investigation which has attracted many workers is that which pertains to the life of the cell—the science of cytology. Although the celltheory was established as early as 1839, little advance was made in this subject in America before 1880. Since that time, however, Americans have been so successful in cytological discoveries that they are now among the world’s leaders in this field.

These studies have been followed along both descriptive and experimental lines. The most prominent of the early workers in this field are E. L. Mark and E. B. Wilson. Mark’s description of the maturation, fecundation, and segmentation of the egg is the most accurate and complete of the early cytological studies. Wilson’s discoveries concerning the details of fertilization and his “Atlas of Fertilization and Karyokinesis,” published in 1895, have now become classic. Wilson, too, has published the only American text-book on cytology,[182] and has more recently taken the lead in studies concerning the relation between the chromosomes and sex. Besides Wilson, Montgomery, Mark, McClung, Morgan, Miss Stevens, Conklin and their associates and students have now furnished conclusive evidence that the sex of an organism is determined by, or associated with, the nuclear constitution of the fertilized egg. This constitution is moreover shown to be dependent upon the chromosomes received from the germ cells.

This explanation is in strict accordance with the results of experimental breeding. It is also quite in harmony with the Mendelian law of inheritance, and in fact forms one of the strongest supports for the view that all Mendelian factors are resident in the chromosomes. Recent work has also discovered the mechanism which governs the complicated conditions of sex which occur in those animals which exhibit alternating sexual and parthenogenetic generations. These remarkable processes are in all cases found to depend upon a definite distribution of the chromosomes.

Other recent experimental work has shown that while the sex is thus normally determined in the fertilized egg, it is in some animals not irrevocably fixed, and the normal effect of the sex chromosomes may be inhibited by abnormal conditions in the developing embryo, as is demonstrated by the recent work of Lillie and others.

The cytological basis for Mendelian inheritance has been very extensively studied by Morgan and his pupils in connection with their work on inheritance in the common fruit fly _Drosophila_. The evidence supports Weismann’s earlier hypothesis that the chromosomes are the bearers of the heritable factors, and that these are arranged in a series in the different chromosomes. This theory is shown to be in such strict accord with both the cytological studies and the results of experimental breeding that Morgan has ventured to indicate definite points in particular chromosomes as the loci of definite heritable factors, or genes.

Confirmation of this view is furnished by the behavior of the so-called sex-linked characters, the genes for which are situated in the same chromosome as that which carries the sex factor. Many ingenious breeding experiments indicate further that all the hereditary characters in _Drosophila_ are borne in four great linkage groups corresponding with the four pairs of chromosomes which the cells of this fly possess.

_Comparative Physiology._

None of the experimental fields has been of greater importance in zoological progress than that which concerns the functions of the various organs. Without this companion science morphology and comparative anatomy would have become unintelligible. American investigators, among whom G. H. Parker stands prominent, have taken a leading part in this field also.

_Neurology._—The physiological analysis of the components of the nervous system, both in vertebrates and invertebrates, is another important branch of experimental biology. The 28 volumes of the Journal of Comparative Neurology attest the large influence that American investigators have had in the development of this science.

_Regeneration._—Experimental studies on the powers of regeneration in plants and animals have been made from the earliest times. During the past few years, however, there has been made a concerted attempt to analyze the factors which determine the amount and rate of regeneration. Much progress has been made toward the postulation of definite laws applicable to the regenerative processes of the parts of each organism. The critical analyses of Morgan, Loeb and Child have been particularly stimulating.

_Tissue Culture._—Another line of experimental work which has been developed within the past few years by Harrison, Carrell, and others is the culture of body tissues in artificial media. These experiments have included the cultivation in tubes or on glass slides of the various tissues of numerous species of animals. They have yielded much information regarding the structure, growth and multiplication of cells, the formation of tissues, and the healing of wounds.

_Transplantation and Grafting._—Closely associated experiments consist in the transplantation of organs or other portions of the body to abnormal positions, to the bodies of other animals of the same species or of other species. In this way much has been learned about the potentiality of organs for self-differentiation, for regulation, for regeneration and for compensatory adaptations. The experiments have shown, further, the independence of soma and germplasm and have revealed the nature of certain organs whose functions were previously obscure.

_Tropisms and Instincts._—Another field of experimental biology concerns the analysis of behavior of organisms in response to various forms of stimuli. These studies are being prosecuted on all groups of organisms, including the larval stages of many animals, and are yielding most remarkable results. The success in this field of research is largely due to stimulating influence of Jacques Loeb, Parker, Jennings, and their co-workers.

_Biological Chemistry._—Still another experimental field which has developed into one of the most important of the biological sciences relates to the fundamental chemical and physical changes which underlie all organic phenomena. A knowledge of both physiological and physical chemistry is to-day essential for all advanced biological work. The peculiar nature of life itself, of growth, disease, old-age, degeneration, death and dissolution are presumably only manifestations of chemical and physical laws. The ultimate goal of all experimental biology, therefore, will be reached only when the basic physico-chemical properties of life are understood. At that time only will the perennial controversy between vitalism and mechanism be ended.

_Economic Zoology._

A moment’s reflection will show that economic biology is the most essential of all sciences to the human welfare and progress. For man’s relation to his environment is such that the penalty for ignorance or neglect of the biological principles involved in the struggle for existence quickly overwhelms him with a horde of parasites or other enemies.

It is only by the intelligent application of biological knowledge that our food supplies, our forests, our domesticated animals and our bodies can be protected from the ever ravenous organisms which surround us.

The losses to food supplies and other products by insects alone amounts to 100 millions of dollars a month in the United States. And the parasites cause losses in sickness and premature deaths each year of many millions more. Then there are the destructive rodents and other animals which add largely to our burdens of support. These enemies next to wars and fungi are the most destructive agencies on earth. Could they but be eliminated man’s struggle against opposing forces would be in large measure overcome. The results of recent work in economic zoology, both in regard to the destruction of enemies and protection of useful mammals, birds and fishes, furnish a bright outlook for the future.

_Protozoology._—Partly as an experimental field for the solution of general biological problems and partly because of its practical applications the study of protozoa has now developed into a special science.

The results of the investigations of Calkins, Woodruff, Jennings and others have greatly supplemented our understanding of the signification of such important biological phenomena as reproduction, sexual differentiation, conjugation, tropisms, and metabolism.

From an economic standpoint the protozoa have recently been shown to be of the greatest importance because of the human and animal diseases for which they are responsible.

_Parasitology._—The animal parasites of man, domesticated animals and plants include numerous species of protozoa, worms, and insects. Together with the bacteria and a few higher fungi they cause all communicable diseases. When we consider that not only our health but also our entire food supply is dependent upon the elimination of these organisms we must admit that parasitology is the most important economically of all the sciences.

The reports of the investigations of Stiles and his associates in the Hygienic Laboratory and of Ransom and his staff in the Bureau of Animal Industry are widely distributed by the federal government. The systematic studies so ably begun by Joseph Leidy in the middle of the last century have been continued by Ward, Linton, Pratt, Curtis and others on the parasites of many groups of animals.

_Economic Entomology._—Another extremely important biological science, the practical applications of which are second only to those of parasitology in importance, is entomology. In the last few years economic entomology has exceeded any of the other branches of biology in the number of its investigators. The American Association of Economic Entomologists has a membership of about five hundred. The work of most of these is supported by appropriations from the State and federal governments, and the results of their investigations are widely published.

It is now well known that some of the protozoon parasites are conveyed from man to man only through the bites of insects. The local eradication of several of our most fatal diseases has recently been brought about by the application of measures to destroy such insects. This is the greatest triumph of economic zoology.

_Economic Ichthyology._—The U. S. Fish Commission has for many years been actively engaged in investigations on the food fishes, including methods for increasing the food supply by suitable protection and artificial propagation. The work includes also edible and otherwise useful mollusks and crustacea. Their marine and fresh-water laboratories have also been of great service to general biological science.

_Economic Ornithology and Mammalogy._—In addition to the local bird clubs and the American Ornithologists Union for the study and preservation of bird and mammal life, the Bureau of Biological Survey has for some years conducted investigations on the economic importance of the various species. The publications of this Bureau are of great value both in determining the economic status of our birds and mammals, and also in recommending means for the protection of the beneficial species and the destruction of the injurious. Several of the States issue similar publications.

_Genetics._

One of the most interesting chapters in biology relates to the development of the modern science of heredity, or genetics.

Previous to the year 1900, when the Mendelian principle of inheritance was re-discovered, the relative importance of heredity and of environment in the development of an organism was little understood. It is true that Weismann had insisted on the independence of soma and germplasm some years earlier (1883), but the body of the individual was still generally considered the key to its inheritance.

The recognition of the general application of Mendel’s discovery gave a great impetus to experimental breeding both in plants and animals. While heretofore it had been necessary to depend upon the somatic characters as evidence of the hereditary constitution of an individual, it now became possible, knowing the hereditary constitution of the parents of any pair of individuals, to predict with almost mathematical certainty the characters of their possible offspring.

In general, the laws of possible chance combinations of any group of characters determine the probability of any particular offspring possessing one or many of those characters. The physical basis for such Mendelian inheritance is evidently the chance combinations of chromosomes which result from the processes of maturation and union of the germ cells.

Certain limitations to the law are met with because the relatively small number of chromosomes involves linkage of genes, because of the occasional interchange of groups of genes between homologous chromosomes, and because the relative activity or potency of any particular gene may differ in different races, and, finally, because the normal activity of any given gene may be modified or inhibited by the action of other genes. It is by no means certain, however, that all inheritance is Mendelian, for there still remains much evidence that the hereditary basis of certain characters may be resident in the cytoplasm, rather than in the chromosomes. A recent book by Morgan, Sturtevant, Müller and Bridges (1915), entitled “the mechanism of Mendelian heredity” gives the cytological explanation of Mendelian inheritance.

Americans have from the first taken a leading part in this field of research and have been quick to recognize its practical applications to the improvement of breeds in both animals and plants. This prominent position is largely due to the experimental work of Castle, Davenport, Morgan, Jennings, Pearl, and their co-workers on animals and that of East, Emerson, Davis, Hayes and Shull on plants.