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FRANKLIN SHULL, Consulting Editor





Metcalf and Flint






There are also the related series of McGraw-Hill Publications in the Botanical Sciences, of which Edmund W. Sinnott is Consulting Editor, and in the Agricultural Sciences, of which Leon J. Cole is Consulting Editor.





Professor of Botany in the University of California at /.> A'tiydcs





COPYRIGHT, 1928, 1932, 1940, BY THE



All rights reserved. This book, or may not be reproduced
the publishers.

parts thereof,

in any form without permission of

The present edition, like the two whicj} have preceded it, aims to present the basic principles common tojill living things, with emphasis on those aspects of biology that seem to be of greatest
value in contributing to a liberal education. Although the general plan of organization remains unchanged, the addition of new material has both broadened the scope of the book and

rendered more thorough the treatment larly those dealing with the animal side This book is intended to serve as a course to give a broad perspective to


many topics, particuof biology. basis for an orientation

the vast field of

objectives arc cultural rather than technical, in biology. of the fact that the great majority of students who recognition elect the elementary course do not later specialize in biology.


the other hand, those who do will be provided with a comprehensive view of the science as a whole and with a foundation of knowledge upon which more advanced studies may be based. The book stresses the " unity of life" concept that, in response to the same natural laws, all plants and animals possess funda-

mental similarity in bodily organization and in vital processes. Elaborate descriptive details have been largely omitted, the emphasis throughout being placed on fundamental facts and

Over one-half of the book deals with the structure, functions, and classification of both plants and animals, while the remaining portion takes up in an elementary way the more general phases of After developing at the heredity, adaptation, and evolution. outset a number of important concepts and generalizations by a discussion of the cell and of unicellular organisms, four chapters
are dbvoted to the plant kingdom. A survey is made of the lower plant groups, and then the seed plants are considered in greater detail, with emphasis on vegetative structure and Next comes an account of vegetative functions reproduction. and irritability. The succeeding seven chapters deal entirely




the important invertebrate

phyla are

material relating directly to the human body has been Vitamins and hormones are dealt with more fully than heretofore. Approximately one-quarter . mussel. 80 new cuts having been made for this edition. This renders the inclusion of a The index. Many changes have been made in the illustrations. metabolic. has been added to a later chapter. two on adaptation. This feature gives the book greater flexibility in permitting the teacher to change the order of topics and to omit material freely without breaking the con- achieve this result. contains all the technical terms used in the text. as related to man. which has been carefully superfluous. starfish. so far as possible. as well as animal tissues. is based largely on the The ensuing discussion of vertebrates Then follow chapters on animal PREFACE earthworm. presented in an ascending sequence. Minor changes have been made throughout the entire book. and reproduction and development. and grasshopper serving as types for the hydra. and in both chapters considerable new introduced. prepared. but it is felt that this is of no disadvantage to the student. The last part of the book includes three chapters on heredity. Technical terms are reduced to a minimum. tissues. a slight amount of repetition has been necessary. functions. The subject matter is presented tinuity of the whole. in a manner simple enough to be easily understood by students in their first year of college work and without any previous knowledge of the subject. coordination. and four on evolution. The author's Laboratory Directions for General Biology" has been revised to correspond with changes made in the present volume and will be found useful as an adjunct to it. the subject .of coordination in animals has been expanded into a chapter separate from the one dealing with their nutritive functions. their respective groups. An effort has been made. it is highly desirable that laboratory work be carried on in connection with a study of this textbook. crayfish. glossary Although not absolutely necessary. The topic of immunity to disease. The first time a term is used. arid definitions are easily found. it is italicized and To either defined or explained. In the present edition. for it is only in the laboratory that a student can obtain " a first-hand knowledge of the subject. in many chapters on almost every page. to develop each chapter as an independent unit.

213. Schwarz for making drawings for Figs. 106. 23(7. 132. and the 26 uncredited photographs. 105. and 209. 40. 206. is grateful to his colleague. Acknowledgment of those reproduced from duplicate electrotypes are indicated by "from". were prepared by the author. Suggestions and criticisms will be welcome. and 114A. of first edition. April. 75. Walthor B. 47. 1097?. 84. about one-half have since been replaced. 196. The author wishes to express his appreciation of the interest by the widespread adoption the former editions have trusts that the book in its present form may be and enjoyed received as cordially. The author is also thankful to Mr. 133. 897?. 1940. 127. 117. 134. while. all of Figs. 79. 26. 109 A and 114B. 163. 28. indicated A. Professor Arthur M. 58. W. 202. tions and for kindly making the drawings for Figs. H. 72. 118. 215. Stebbins for Figs. 124. 225. were kindly made by Miss Alice Handsehiegl: for the following figures. All the other original drawings. The author 43. 197. 204. 86. in the preparation of many of the illustraassistance for Johnson. 108. and 234. 119. 217. 208. Robert C.PREFACE of the old figures vii have been replaced by new ones." kindly granted permission to use illustrations from copyrighted sources. 200. All the illustrations not credited to others are original. 198. The total number of illustrations has been increased over the second edition by 11 and over the first edition those appearing in the by 32. numbering 90. 205. also to Mr. 67. 49. UNIVERSITY OF CALIFORNIA AT Los ANGELES. those redrawn from other books by Thanks arc due the publishers and authors who have "after. which appeared in the Drawings first edition. 70. 71. 216. 61. . 894. 207. 214. 56.


. monocotyledons ture. tissues. CHAPTER V VEGETATIVE ORGANS OF SEED PLANTS 57 Structure of root tip THE ROOT Organization of root systems of leaves Structure of mature root Arrangement ture. . Food storage in roots THE STEM Buds and branches. PAGE v CHAPTER INTRODUCTION Features I 1 common to plants and animals II Scope of biology Applications of biology. and generalized cell systems Structure of protoplasm Composition of protoplasm A Behavior of protoplasm Nature of life Spontaneous generation The cell principle. CHAPTER UNICELLULAR ORGANISMS Protococcus cella ITI 18 - Yeast Euglena Bacteria Amoeba Colony formation . CHAPTER PROTOPLASM AND THE CELL 5 Multiplication of cells Tissues. Stems of Young woody Mechanical stem. THE LEAF External features Internal struc- CHAPTER REPRODUCTION IN SEED PLANTS VI 82 THE FLOWER Pollen and Pollination Ovule and embryo sac Floral parts pollen tube Fertilization Alternation of generations ix .CONTENTS PREFACE.-Habit Stem strucIncrease in diameter. Conducting Herbs and woody vines. tissues. . organs. . Paramccium Vort i - CHAPTER THE LOWER PLANT GROUPS THALLOPHYTES TV 35 Asexual reproduction Sexual reproduction Groups of algae FUNGI Vegetative body Reproduction BRYOPHYTES Vegetative body Sexual organs Sporophyte Alternation of generations PTERIDOPHYTES Vegetative body ALGAE Vegetative body Spore production Gametophyte Heterospory.

Digestive system. Excretory system. Reproductive Development THE GROUPS OF VERTEBRATES Fishes Amphibians Reptiles Birds Mammals. Circulation in mammals. CHAPTER VII METABOLISM AND IRRITABILITY IN PLANTS 100 Independent and dependent plants Absorption Diffusion Osmosis Plasmolysis Conduction and transpiration PhotoFormation of fats and proteins Utilization of food synthesis Food storage Digestion and assimilation Respiration Irritability. Other crustaceans The myriapods The grasshopper. Circulation in fishes. . Nervous system. Excretory system. Circulatory system. Reproductive system Other insects The arachnids General features of arthropods. Reproduction CHAPTER X THE VERTEBRATES CHORD AT A The worm. The lancelet. Reproduc- Other echinoderms. Structure. Nervous system. External features. Reproduction system Other ECHINODERMATA The starfish. Skeletal system.x in seed plants tive CONTENTS PAGE features Embryo and endosperm The fruit THE SEED Seed of gyrnnosperms for Reproducstructure of Dormancy seedling Conditions germination Development the VEGETATIVE PROPAGATION. Circulatory system. system. 143 fresh-water mussel. CHAPTER IX THE HIGHER INVERTEBRATES MOLLUSCA cavity. Respiratory system. External features. Nervous system. Circulation in amphibians. Excretory system. External features. CHAPTER XI CHIEF ANIMAL TISSUES Epithelium 190 Muscle tissue Nerve tissue Connective and sup- porting tissues. Reproduction COELENTE RAT A The hydra. structure. reproduction Other coelenterates PLATYHELMINTHES NEMATIIELMINTHES ANNELIDA The earthworm. CHAPTER THE LOWER ANIMAL GROUPS VIII 118 PROTOZOA PORIFERA Structure. Internal anatomy. External features. the sea 163 squirts pro vertebrates. Digestive system. Respiratory system. General annelids tion features. Digestive system. Circulatory system. Mantle and development Other mollusks Reproduction The The appendages. The acorn-tongue The frog. ARTHROPODA The crayfish.

Cursorial adaptation. CHAPTER XIV REPRODUCTION AND DEVELOPMENT IN ANIMALS Asexual reproduction Sexual reproduction Germ cells Fertilization Parthenogenesis Early embryonic stages Gastrulation Mesoderm and coelom formation Later development Oviparity and viviparity Intrauterinc development Larval stages Parental care. Significance of alternating generations Determination CHAPTER XVI MENDELIAN LAWS OF HEREDITY Hereditary elements Gene combinaPrinciple of segregation Backcrosses Dominant and recessive characters Printions ciple of free 261 assortment Linkage Sex-linked inheritance. The eye Chemical coordination.CONTENTS xi PAGK CHAPTER METABOLISM IN ANIMALS Feeding habits XII 197 Dietary requirements Vitamins Digestion Human digestive system Digestion in man Absorption Respiration Circulation Excretion. Arboreal adaptation. Natatorial adaptation. CHAPTER XVII APPLICATION OF HEREDITARY PRINCIPLES 280 BREEDING Variation Mass culture Pedigree culture Hybridization EUGENICS Heredity and environment Hereditary characters in man Aims of eugenics PLANT AND ANIMAL Elimination of defectives Increase of superiors. CHAPTER XIII 215 COORDINATION IN ANIMALS Nervous system of man Neurons arid nerves Reflex action The spinal nerves Sense organs. Fossorial adaptation. The ear. Volant adaptation Adaptive radiation . CHAPTER ADAPTATION RACIAL XVIII 290 ADAPTATION pollination Insect Desert dispersal plants Seed Protective resemblance Feet and bills of birds Generalized mammals Teeth of mammals Limbs of mammals. 228 JCHAPTER XV PHYSICAL BASIS OF HEREDITY Uniparental inheritance Biparental inheritance The hereditary bridge Vegetative or somatic mitosis Significance of fertilization 245 Reduction of chromosomes Grouping of chromosomes in gametes of sex.

ing bacteria CHAPTER XX THE FACTS OF EVOLUTION Importance of evolution evidence Classification 335 Popular misconceptions Vestigial Nature of the In- structures Embryology tergrading species distribution. Malarial paraImmunity SYMBIOSIS Lichens Nitrogen-fixsite. PARASITISM. Tapeworms Ants arid Aphids Hermit crabs and sea anemones. Adaptive re- CHAPTER XIX 317 SAPROPHYTISM. CHAPTER XXII THE CAUSES OF EVOLUTION INHERITANCE OF ACQUIRED CHARACTERS Direct action of environment Use and disuse NATURAL SELECTION Overproduction Competition variations -Variation 389 Survival of the fittest Limitations of natural selection MUTATION Continuous and discontinuous CLUSIONS to natural selection Relation Objections Causative factors Directive factors.xii CONTENTS PAGB Convergent adaptation INDIVIDUAL ADAPTATIONS sponse Explanation of adaptation. AND SYMBIOSIS SAPROPHYTISM Decay of organic mat tor -Cycle of elements PARASITISM Parasitic plan in Parasitic animals. Cultivation and domestication Geographical CHAPTER XXI THE LIFE OF THE PAST Nature and formation of Length of geologic time fossils 351 Division of geologic time of* the THE ARCHEOZOIC AND PROTEROZOIC Life ERAS Life of the Archeozoic Proterozoic Early THE PALEOZOIC ERA Cambrian stages in evolution vician Carboniferous and Silurian and Devonian and OrdoPermian THE MESOZOIC ERA Life of the Cretaceous Life of the Triassic Life of the Jurassic THE CENOZOIC ERA Life of the Tertiary Life of the Quaternary EVOLUTION OF THE HORSE AND ELEPHANT Adaptive features of modern horses Stages in equine evolution Adaptive features of modern elephants Stages in evolution of the elephant CONCLUSIONS. 425 427 . CON- CHAPTER XXIII THE EVOLUTION OF MAN Distinctive features of primates The anthropoid apes Distinctive features of man History of the primates The Java ape 404 man man APPENDIX INDEX The Peking man The Piltdown man The Heidelberg The Neanderthal man The Cro-Magnon man.

zoology. processes. based on a study of the higher members of each kingdom. This misconception arises 1 from the occurrence of certain .FUNDAMENTALS OF BIOLOGY CHAPTER I INTRODUCTION all In its broadest sense. Because of the occurrence of organisms intermediate in certain respects between typical plants and typical animals. typ. yet nearly all of them can be referred to either the plant or the animal kingdom. the separation of all living things into these two conventional groups is often an uncertain matter. all complex. arid laws peculiar to them are said to be organic. and higher forms of and plants animals. animals. Distinctions between plants and animals.e of organization differs in the lower Nevertheless. not always easy to distinguish between the two forms in which life commonly exists.eial study of and called of plants. are known as organisms. Thus the spe. and other lifeless things. it is they are called organisms. The multitudinous forms of life inhabiting the earth are infinitely diverse in regard to structural details and life habits. are to be subdivisions of the the two as major larger science of regarded biology. the facts arid principles that have been derived from a scientific study of living things. are rather easily made. and all substances. it is no exaggeration to say that all It is life is one. but when lower forms of life are examined. called botany. And because even the most highly differentiated plants and animals have many fundamental features in common. For The life. it includes Biology is the science of life. Plants and animals possess a characteristic bodily organization that at once sets them off from minerals. rocks. whether simple or this reason Although the animate is sharply set off from the inanimate. commonly supposed by many people who have not studied biology that plants and animals do not manifest the same kind of life. these distinctions break down.

The same natural laws apply to all organisms. is the most fundamental division of biology. Moreover. and oysters do not. (6) The essential features of reproduction (7) are common to plants and animals. distinctions are superficial state of living. assimilation. In fact. such as locomotion and Although most plants are stationary and are slow sensitivity. collectively known as metabolism. Naturally. barnacles. is common to all living things. some plants have sensitive leaves or floral organs that react to stimuli than do such sluggish animals as sponges. is too vast a subject to be mastered in its entirety by any one man. These are as follows: (1) Morphology. (2) This living matter is organized in both plants animals into microscopic units called cells. called protoplasm. These processes include digestion. These may be stated as follows: (1) Life is always associated with a unique substance. to respond to external influences. corals. absorption. composition. which in all plants and animals is essentially similar in structure. The distinction between living and non-living things rests upon certain basic similarities in organization and behavior shared by all organisms. it is possible to consider. such as the laws of heredity and evolution. much more quickly Features Common to Plants and Animals. (4) The property of irritability. consideration of gross features (anatomy) as well as the minute . and (3) Certain vital processes. the study of the form and structure of organ- It includes a isms. in an elementary course in general biology. This has been made necessary for purposes of advanced study because biology.2 FUNDAMENTALS OF BIOLOGY conspicuous differences between them. and behavior. Living things may be studied from many different aspects. take place in plant bodies in essentially the same manner as in animal bodies. which is the power of responding to external influences. most animals move freely from But these place to place and exhibit a high degree of sensitivity. each of which has been organized as a special branch of biology. (5) Growth in all many-celled organisms takes place by a complicated process of cell division followed by cell enlargement and cell differentiation. Scope of Biology. and respiration. only the more basic subdivisions of the subject. in a broad way. while such animals as sponges. and have nothing to do with the actual some of the lower plants have the power of locomotion. including all organized knowledge pertaining to living things.

(3) Taxonomy is concerned with the of and classification naming organisms and represents the oldest branch of biology. the English sparrow. It is Organic evolution is a study of the concerned with the history of life . temperaAll living things exhibit adaptation to the conditions ture. Solanum tuberosum. moisture. 1707-1778. Every Fio. Ecology is a newer field of biology that takes up the life relatheir relations to tions of organisms one another and to various factors of their environment. in terms physiology. by the great Swedish naturalist. so far as possible. etc. of physics and chemistry. Passer domesticus. the dog. 1. the white oak is Quercus alba. known species of plant and animal has been given a scientific name consisting of two parts.INTRODUCTION 3 details that are seen only with the aid of a microscope (histology) with vital processes and (2) Physiology deals with functions . and reproduction belongs to the field of These it seeks to explain. growth. Plants and animals are named according to a binomial system devised von Linne*. A study of the functions concerned with metabolism. which indicates natural relationship. Organisms are classified into groups on the basis of fundamental structural (4) resemblance. Carl more commonly known as Linnaeus (Fig. Carl von Linn6. Canisfamiliaris. under which they live. such as light. For example. the common potato. activities. irritability. (5) descent of organisms. 1).

man's principal Scientific knowledge. home economics. (2) Biology is a necessary There are many special fields of prerequisite to further studies. dietetics. or to some extent. such as wool and silk. From a purely (1) It is pursued for intellectual gratification. while serums. horticulture. etc. . hygiene. Familiarity with. An elementary study of biology is given a prominent place in college curricula for several reasons. Wood has always been a building material of first importance. cultural standpoint biology is of considerable value in giving one an acquaintance with the world of living things. and many others. living In fact. contributing enormously to his welfare they and comfort. make human either a plant or an animal from both plant fibers. agriculture. Many medicines are derived from plants. the human body comes from life possible.. (4) Plants and animals arc of inestim- able material value to man. studies of plants and animals.4 FUNDAMENTALS OF BIOLOGY on the earth and with the structural changes that the various existing species have undergone in the course of their racial development. are organic in origin. It is concerned with the resemblances and differences between individuals. sanitation. Applications of Biology. These include medicine. an appreciation of the phenomena of life. knowledge that are based largely. or participation in. (6) Genetics is a new field that has grown out of the study of evolution. an elementary knowledge of biology gives one a basis for an understanding of his own body and thus is a direct aid to health. especially those due to heredity. cotton and linen. based on fuels. From All the nourishment that Man makes his clothing utilization of many of these products. coal. and an understanding of some of the and processes of nature. and petroleum. the those same to laws as governing all living organism subject is built the structural and to same according plan as other things mentary training in highly developed animals. is essential to the most efficient enters things man derives his food. forestry. psychology. on biological great laws facts and principles. such as source. soci- ology. vaccines. Wood. and animal fibers. come from animals. any of these fields requires at least an ele- (3) Because man is an general biology.

seen in Figs. 2 and 3. where several kinds of simple cells are in plants posed protoplasm. These are known as cells. These may be readily all have certain basic features in common. This is deposited by the protoplasm 5 . the cytoplasm by a plasma membrane. the nucleus. 2 and 3). This means that all vital activities the processes that go on in plants and animals. Recognition of the fact that cell is the unit of structure and function is fundamental to an understanding of much of the subject matter of biology. is nuclear material being. more complex. Cells ai^e not only units of structure but are also units all of function. a starfish's egg. The nucleus enclosed by a delicate nuclear membrane. a moss leaf. a thicker covering composed of an organic substance called cellulose. and so it is appropriate to begin our studies of organisms with a consideration of the cell and of the living matter of which it is com- the Although there are many kinds of cells and animals. Almost all plant cells have. A Generalized Cell.CHAPTER II PROTOPLASM AND THE CELL When is examined with a microscope a small isolated portion of an ordinary plant or animal it is seen to be composed of a . each but represents living matter of a different kind. the toplasm. which differ considerably in many ways. or any other favorable material (Figs. A typical cell consists essentially of a dense spherical body. organized masses of living matter. surrounded by a mass of less dense material Both the nucleus and cytoplasm consist of procalled cytoplasm. Simple types of cells may readily be seen when we look at a piece of a salamander's skin or a drop of its blood. a thin section of a rootlet. in general. Extensive microscopic examination of the most diverse kinds of plants and animals has demonstrated that cells are the units of which living things are constructed. in addition to the plasma membrane. shown. just as individual stones or bricks may be the components of a wall. great many minute.are performed by various kinds of cells.

while older of cells generally assume a variety of shapes. pm. cell from a hair r . young colls. cyt. although eye. This is particularly true of. Some X 350. cytoplasm. nearly all vary widely in size. FIG. them are too small to be seen with the naked . X 750. X 150. B. this For the most part their cells lack cell walls. although not living. E. chl. in plants. on the contrary. cw. often Cells also becoming lengthened. nucleus. In animals. constituting the cell wall. section of an unfertilized starfish egg. but cells in contact with others may be flattened along the sides by mutual pressure. 2. each mass of protoplasm being separated from adjacent ones merely by its thin. giving them a polyhedral form. Because. cell wall. Free cells tend to be spherical. red blood cells of a salamander. C piece of the outer skin of a salamander. nuc. the the protoplasmic units arc separated from one another by partitions of non-living substance. generalized plant and animal cells. living plasma membrane. the living matter itself occupies little rigid box-like is not the case. is considered part of cell. of a squash leaf. vacuole. X 500. X 250. nuc A. vac. compartments. cells from a moss leaf. plasma membrane. D. chloroplast.6 that it FUNDAMENTALS OF BIOLOGY encloses and.

3. which are filled with a fluid designated merely as Vacuoles are generally larger and consequently more In fact. older cells. spherical bodies known as nudeoli are also present within the nucleus. mature cell. and less commonly in animal cells. cell The cytoplasm is finely granular and ordinarily less dense than The minute granules may surround clear spaces sap.000. and plays the principal part It is of great in the process of cell importance in connection with heredity. The cytoplasm of very young plant cells is relatively dense and contains many small vacuoles (Fig. showing tho enlargement of vacuoles which accompanies growth of the cell. 3). in the latter striking in plant cells than in animal cells. As the cell enlarges. the nucleus being suspended in the center of the cell by cytoplasmic strands. the nucleus. A FIG. the cytoplasm often appears essentially homogeneous. Finally there is often just . the vacuoles coalesce. called vacuoles. Every typical of a fluid. A. in which is embedded a coarsely granular substance called chromatin. which consists principally the nuclear sap. Generally in plant cells. X 1. one or more small. The nucleus seems to control the metabolic activities of the cell division.PROTOPLASM AND THE CELL cell contains a nucleus. C. dense. Cells B C from a longitudinal section of an onion root tip. B. young cells.

photosynthesis. Tissues. examples of tissues made up of highly specialized cells. blood vessels. A system is a group of with concerned some For organs general bodily function. in turn. the heart. while tissues. brain. and nerve and muscle in animals are functions. Cells exhibit the striking peculiarity Multiplication of Cells. etc. cells results in the formation of tissues. this is a and animals in structure. contain wall. cells All are generalized. a ing in cell volume by the assimilation division the protoplasm in each daughter cell increases of food and the absorption of water. or a heart.8 FUNDAMENTALS OF BIOLOGY one large vacuole around which the granular portion of the cytoplasm occurs as a thin layer. Chloroplasts are dense masses of protoplasm colored by the green pigment They carry on the very important function of chlorophyll. the nucleus lying against the cell Cells from the green parts of plants. composition. alive. Followdivision. or stomach. while embryonic cells and a few mature most mature cells are specialized for particular functions. of being able to increase in number and thus to reproduce themThe formation of new cells is accomplished by cell selves. and blood make up the circulatory system. and In view of the enormous diversity among organisms. as from a leaf. Structure of Protoplasm. A tissue is a group of similarly differentiated cells performing one or more particular Wood in plants. the only substance that similar in all plants by no means is identical. and consequently are structurally differSuch a " division of labor" among various groups of entiated. such as a root or leaf. liver. cell enlargement. Cells that do not carry on a definite kind of work are most nearly like the generalized cell thus resulting in that has been described. An organ is merely a differentiated part of the body. and Systems. example. Organs. and other organs of digestion comprise the digestive system. remarkable . and spherical or discoid (Fig. generally are grouped to form organs. Cells are organized as tissues in all plants and animals except the lowest. specialized bodies called chloroplasts. Although protoplasm. These are generally small. complex process involving first a division of the nucleus and then a separation of the cytoplasm into two parts. whereby the plant is able to manufacture food under the influence of light. fact. the stomach. numerous. is essentially behavior. 2 A).

The liquid inside the vacuoles differs in nature properly fixed and stained. is nearly always distinctly granular in appearnoted. examples being gelatin. the dispersion medium is water containing salts a suspension. Matter in the colloidal state consists of finely divided particles dispersed through a A continuous medium. however.PROTOPLASM AND THE CELL 9 fluid substance. appearance similar to that protoplasm ordinarily presents an from that surrounding them. and other dissolved substances. Protoplasmic structure can be simulated by adding a small quantity of lampblack to an emulsion of olive oil and water. which is fluid. The carbon particles represent the oil globules the vacuoles. Colloidal particles. and is also almost transparent. are sometimes large enough to be seen under the microscope. methods of fixation and staining have been devised so that the structure of a viscous liquid. but are smaller than the particles in In the case of the colloidal substances present in protoplasm. Living matter. it usually has a As already slightly grayish tinge. The great variety of reactions taking place doubtless arises from the tremendous . gum arabic. the distributed granules being throughout a clear liquid or gelatinous medium which surrounds globules of various sizes. and the the protoplasmic granules. as the particles in a true solution. protoplasm may be seen more clearly. as by a change in temperature. but are usually ultramicroscopic. to the other can be brought about readily by external conditions. which is semisolid. the globules being vacuoles. mixture may exist either as a sol. and egg albumen. The consistency of protoplasm varies from the sol to the gel state. a change from one state as a gel. but is generally more liquid than solid. They are not so small. protoplasm ance and usually contains vacuoles of various sizes. as seen under the microscope. On account of its nearly colorless and almost transparent quality. which do not exceed one molecule in size. Whether living or of a fine colloidal emulsion. is usually a semibut varies from a jelly-like consistency to that of Although nearly colorless. The colloidal nature of the chief constituents of protoplasm the proteins has an important influence on the colloidal activities that A go on within living cells. generally representing aggregations of molecules. or As a rule. water the medium in which the granules are suspended. its fact of considerable importance regarding protoplasm is that constituents form a colloidal system.

contain the four elements. and fats will now be briefly considered. 2. organic substances many different kinds. are very common in non-living matter.0 per cent. Other inorganic constituents of protoplasm are certain dissolved gases. 65. Without this large proportion of water. The 12 elements mentioned above (with the exception of oxygen. Composition of Protoplasm. and chlorine. and even to the same cell at consecutive Yet numerous analyses have shown that periods of time. oxygen. carbohydrates. The most abundant and most important in protoplasm are proteins of egg. approximately 97 per cent of ordinary protoplasm consists of four elements that occur. potassium.5 per cent. but a mixture of many different kinds of substances. Gases and salts occur only in small sulphates. as "living substance. nitrates. and carbonates. such as oxygen and carbon dioxide." it Although commonly spoken of is must be understood that protoplasm not a single substance. that these is made up iron. sodium. protoplasm contains only a relatively few elements. The most important compounds water. quantities.5 per cent. and dissolved salts. Only rarely are It is important to realize that traces of other elements present. This variability extends to different parts of the same organism. on ijhe average. salts. White of and the gluten of cereals are examples of sub- Proteins always stances almost entirely protein in composition. samples from different organisms being essentially similar. commonly forming 85 to 90 per cent of its weight. nitrogen. The remaining 3 per cent 11. lean meat. phosphates. The most abundant constituent of active protoplasm is water. and . of minute amounts of sulphur. both organic and inorganic. complex. hydrogen. some of which are chemically simple. and that protoplasm contains no elements distinctive of itself. magnesium.0 per cent. carbon. and of these there are a great many kinds. which may occur free as well as in combination) always occur in living matter in the form of compounds. 18. living matter cannot carry on its ordinary functions. hydrogen. proteins. others very Protoplasm is somewhat variable in composition.10 FUNDAMENTALS OF BIOLOGY* complexity and instability of the heterogeneous colloidal system that protoplasm represents. calcium. in about the following proportions: oxygen. carbon. chiefly chlorides. but never exactly alike. phosphorus.

it possible for what we call life to express itself. In fact. for example. Proteins of therrTsatisfactorily. life may be merely a result of the organization itself. Behavior of Protoplasm. but usually sulphur 11 and sometimes phosphorus are Protein molecules are very large and extremely present also. but exhibits a very ties in . In addition to the proteins. This means that its constituents are physically related to one another in ways that are extremely intricate. Examples of carbohydrates are sugars. hydrogen. and cellulose. realized that all our knowledge of its composition has been (2) The composition constantly fluctuating as a result of chemical changes that go on all the time. it is apparent that we can know what substances are present in a given cell only at the particular moment when analyzed. has been roughly determined as follows: CcgeHimNmC^oSs. When liquid. starch. and thus its composition may be altered. Protoplasm is not only the most complex material in existence. and oxygen. In fact.PROTOPLASM AND THE CELL nitrogen. and that therefore It is this organization that makes living matter is a system. being made up of hundreds or even thousands of atoms. protoplasm usually contains other up from simpler compounds of compounds. (3) Protoplasm is not merely a collection of various substances. however. both of these substances are composed entirely of carbon. Unlike the proteins. that they are built called amino acids. the composition has been which accurately determined in many cases. complex. of living matter is of this fact. among which should be mentioned the carbohydrates and fats. The are so complex that chemists have never been able to analyze any It is known. It is not difficult to distinguish between animate and inanimate things because we recognize the presence of life by certain sharply defined criteria by peculiarithe behavior of living matter. they are termed oils. Fats differ from carbohydrates in containing less oxygen in organic proportion to the hydrogen present. empirical formula of egg albumen. foregoing facts regarding the chemical composition of protoplasm must be accepted with three reservations: (1) When The subjected to chemical analysis. In view derived from a study of dead protoplasm. the instability of protoplasm is one of its most characteristic attributes. protoplasm must necessarily be It should be killed. but is undoubtedly an organization of substances.

In some cases. called the "physical basis of life. especially in Paramecium and in certain plant cells. Life without respiration is impossible. but by the interposing of the new 3. In some cases protoplasmic movement results in locomotion. a decomposition process involving the absorption of oxygen. chemicals.12 FUNDAMENTALS OF BIOLOGY These are manifested remarkable and unique set of properties. Nature of Life. respectively. does not move contact. which is the capacity of responding to such external influences as gravity. so that they It is the property of of the living organization. This occurs. Protoplasm. not by the addition of layers on the outside (accretion). Protoplasm exhibits irritability. light. amoeboid and ciliary movement. its particles among become part 4. supposed from causes inherent in the living matter itself. assimilation that makes growth and reproduction carries possible. When stimulated by these influences. Although the preceding criteria enable us to distinguish living from dead objects. as follows: 1. various cells react in characteristic ways. a rotation or streaming of the cytoplasm occurs within the cell boundary. heat. to result Protoplasm has the power of independent motion. 2. regarding the real nature of life. which the famous English zoologist. Irritability is one of the most distinctive features of protoplasmic behavior. but the cell as a whole traction of muscle or change its shape as a result. The con- and the consequent movement of some part of the body depend upon the ability of special contractile cells to change their shape. electricity. Living matter on respiration." is ordinary matter in a peculiar . like the growth of crystals in a supersaturated solution. Thomas Henry Huxley. as a result of which organisms become adjusted to their environment. it is impossible to define life except in its own terms. living matter takes up from environment inanimate substances of varied composition and from them increases its mass. biologists as yet know practically nothing. By the process of assimilation. are characteristic of the onecelled animals Amoeba and Paramecium. called other cases not. two of which. In fact. and is just as characteristic of plants as of animals. and certain others. in Several types occur. the liberation of energy. and the formation of waste products. the old ones (intussusception).

and each year finds new ones being added to the list. One of the strongest arguments is that protoplasm has never been made artificially in the laboratory. when urea was some time it will be. The vitalistic view of life is that protoplasm owns its peculiar behavior to the presence of some kind of special force or " vital that life is something which enters into protoplasm principle" and enables it to function. apparent that the chief point of difference between these two theories is that the mechanists regard life as a result. It maintains that living matter owes its distinctive properties to the little highly complex composition and interaction of the various substances composing it. if true. while of the vitalists the vitalists regard it as a cause. it is probably useless to solve the greatest problem of all time the nature of to attempt For this reason. But since 1828. two theories have been held: the mechanistic theory and the vitalistic theory. among modern It is biologists. there is very difference between animate arid inanimate things. This theory contends that life is merely the expression of certain physical and chemical laws that still are very imperfectly understood. has been to observe. This "vital principle " has never been identified or analyzed and perhaps from its can be. vitalism does not have much support life.PROTOPLASM AND THE CELL state 13 matter with a unique set of properties. According to the mechanistic conception of life. because* very little is known concerning About all the causes underlying the behavior of proto- lias been accomplished in an attempt to plasm. To this the mechanists reply that perhaps all organic the be within bodies could formed of only compounds organisms. Regarding the causes of vital phenomena. for. . scientific Consequently the vitalistic theory is very nature never not conducive to investigation. measure. and record the ways in which they are mani- that fested in plants and animals. It was once thought that synthetically. understand the distinctively vital phenomena." it is only by accepting the mechanistic point of view that progress can be made toward a first made fuller understanding of the nature of life. There is almost total ignorance regarding the nature of life. thousands of different organic substances been have made in chemical laboratories. as their name itself signifies. Thus even though there may be such a thing as a "vital principle.

" Several investigators attempted to disprove the spontaneous origin of these minute life by showing that they enter from the air. but their met with only partial success. and maggots were claimed to come directly from decaying meat. Redi placed three jars of meat in an open window. a world of life was discovered. but no eggs were laid on the B. In 1668. invention of the microscope in the seventeenth century. it to attract the These and other similar experiments did a great deal toward But with the dispelling *the theory of spontaneous generation. He showed that maggots do not arise from decaying meat de novo. but maggots developed only in the jar which had been left uncovered.C. Redi observed flies in in the meat contained the uncovered jar their eggs depositing and on the gauze of the second jar. and it was rather of minute forms If revived. many strange One of ideas have been held in the past concerning their origin. by a series of very forms of efforts careful experiments. great numbers of microorganisms soon appear. conducted a set of experiments in an attempt to disprove spontaneous generation. All the meat decayed. Louis Pasteur (Fig.14 FUNDAMENTALS OF BIOLOGY Spontaneous Generation. even among scientific men. One was left uncovered. demonstrated conclusively that microorgan- . Frogs and toads were supposed of ponds under the influence the from at bottom to develop mud into water were thought to give rise the horsehairs of falling sun. but always develop from the eggs of flics deposited there. which has had many supporters. naturally thought that they arose directly from the decaying organic matter in the "hay infusion. the most widespread theories has been that of spontaneous generation. to worms. This idea was prevalent from the time of Aristotle (384-322 middle of the nineteenth century. It was not until 1864 that the theory was completely overthrown. and the third covered with parchment. Many other similar ideas were common. an Italian. Francesco Redi. Because of the great abundance and rapid ^multiplication of certain forms of life. one covered with gauze. It held that of life can arise directly from inorganic matter low forms many or from dead organic remains. 4). and the theory was a small quantity of hay is boiled and the liquid placed in a covered vessel. because no odors could pass through flies. These were seen with the microscope.) until the parchment. It was then that the great Frenchman.

Nothing came into existence. he boiled liquids containing material capable of yielding jn'n-nx filtrating it :<.:! through cotton. science of bacteriology benefactors. 4.* to the liquids. protoplasm comes only from antecedent protoplasm. that dust particles result of Pasteur's experiments to disprove the theory of spontaneous generation. whatsoever. . his experiments. Omne vivum ex vivo. this proved carry i-ii'TiMi-ti. As far as we can tell. like the nature of life. is no of there scientific the origin of the first forms evidence life. even.i: N:i. As a i-m-. Louis Pasteur. 1822-1895.PROTOPLASM AND THE CELL 15 isms do not appear in sterilized organic matter. he laid the foundations for the great biological and then admitted air to them after Since no decay resulted. Every kind of plant and animal originates from preexisting individuals of its own kind. and life There as In regard to presents an unbroken chain from the beginning. if exposed In one of to air. and became one of mankind's greatest is now no reliable scientific evidence that living things. we know them. ever arise by spontaneous generation. provided that the air is entirely free of dust. FIG. is known as to how the first protoplasm The origin of life.

This is shown by the the word "cell" itself. but their importance was not appreciated for nearly 175 It was then that the "cell years after Hooke's discovery.16 is FUNDAMENTALS OF BIOLOGY may or an unsolved problem. who He perceived with a simple microfirst described them in 1665. however. to include the protoplasm as well. The Cell Principle. Schleidon established the cell kingdom in 1838. may The fact that the cell is the fundamental and animal structure has been recognized only The discovery of cells was made much earlier. the secret of which science not be able to reveal at some future time. and where extended gradually no cell wall is present. made to it. who scope of his of claimed that in the animal all organisms are composed of theory in the plant cells. added. theory" two Germans. Schwann kingdom the following year. Other observers saw and studied cells in many kinds of organisms. that ordinary bottle cork is minute compartments to which he gave the name composed Cork tissue forms the outer bark on old stems. cork wood and example. tant addition has been of the material in the bodies of the higher organisms consists of substances formed by cells but not themselves living. comof cells. being credited to the Englishman. It consists entirely mercial cork coming from a species of oak. for the outer part of the skin. arid Schwann was that the bodies of all plants and all animals To this statement must now be are made up entirely of cells. shells and hair in animals. of dead cells. however. mineral matter in the teeth and bones. on based the was thorough investigations of announced. " it is apparent that much their because "and products. the fluid portion of the blood. as in plants. so that today the gators An imporcell theory is a fully established biological principle. indicating a box-like compart- . etc. Robert Hooke. Although Hooke saw and described the meaning of the term he introduced has been ernpty cells. while the contents were regarded as unessential retention of and were largely ignored. which should be feature of tho cell theory of Schleiden The essential recognized. the protoplasm having completely disappeared and left only the cell walls. The earlier investigators considered the cell wall to be the most important part of the cell. Subsequent investihave fully substantiated their results. Matthias Schleideii and Theodor Schwann. since 1839. the protoplasmic unit itself is called a cell. unit of all plant own manufacture.

established the similarity between the living matter of both plants and animals. the basis of life. to appreciate the importance of the cell contents and to In 1835. however. he applied the term give a name to the living substance. the German botanist. as a the first 17 The Frenchman. Hugo von Mohl. . Since that time the term protoplasm has been applied to all living matter.PROTOPLASM AND THE CELL ment. was cell in a prison. but thought it limited to them and did not regard it as In 1846. Felix Dujardin. It was not until 1861. that the work of Max Schultze. sarcode to the material composing the bodies of microscopic animals. another German. although it had been used earlier in a different sense. gave the name protoplasm to the living contents of plant cells.

living alo^iie. and It can yet a single cell. Protococcus. and the resemblances and differences all of carry on between them noted. Obviously. is one of the simplest forms of plant life (Fig. etc. although doing its work in a much simpler way.. This plant. walls. that a plant or an animal may consist of many cells or of just one The lowest forms of life are unicellular. the nucleus is difficult to see. each individual cell. known as Protococcus viridis. scrape off a bit of this material. representative types of unicellular organisms will now be discussed. independent 1 organism. Several kinds of common. cells have been considered as struccomposing the bodies of many-celled plants and but among the lowest organisms cells can carry on an animals. the limits of which are difficult to make out in the living cell. this represents the simplest possible condition of structural organization. rocks. In the presence central nucleus 1 Often called Pleurococcus vulgaris. Each cell consists of a mass of cytoplasm surrounding a small tree trunks. we see innumerable small. There is often present on the shaded side of If drop of water. In the living condition. a bright green stain.ORGANISMS In the preceding chapter. This means ism. 5).CHAPTER III UNICELLULAR. but in reality is confined to a single large peripheral chloroplast. spherical. is a complete individual. or it may exist as a separate organism in itself. The green coloring matter. the functions that multicellular individuals perform. tural units being a single unit of protoplasm. is a pigment that appears to be uniformly diffused throughout the cytoplasm. independent existence and thus live entirely apart from other Therefore a cell may be merely a minute part of an organcells. mount the scraping in a damp we and enclosed by a cell wall composed of cellulose. 18 .500 inch in diameter. and examine with a microscope. called chlorophyll. green cells averaging about 1/2. Each cell is an individual plant a complete.

cell the is It under influence called of green light photosynthesis. undergoes division into two cells. when The making of sugar in a illuminated. nature. but like all other organisms it also carries on reproduction. substances from which sugar is made. but all green cells. Thus the The sugar made by presence of chlorophyll enables Protococcus to make its own food. New individuals come into existence by a method called fission. X cell 1. These two processes should not be confused. viz. First the cell elongates Then a cell wall forms between slightly and its nucleus divides. and respiration.. Each Some of the contains a nucleus. The inorganic FIG. a simple one-rolled plant. which proceeds only in the daytime and only in green cells. it is assimilated by the living protoplasm. This is in all living cells Protococcus not only performs the strictly metabolic processes photosynthesis. the cell carries on a very important function by It makes sugar the manufacture of sugar. tion. cells. the cell is used as a source of nourishment. After a cell has of reached a certain definite it size. a substance that enables it to carry on respiraan energy-releasing process that goes on constantly and is entirely distinct from photosynthesis. and this ability it shares with all green cells. and a single lohcd chloroplast. can carry on this process. if external conditions are favorable. cells 5. is one of the most important processes in. water and carbon The cell also dioxide. absorbs oxygen. assimilation.000. Not only Protococcus. Protococcus viridis. . that is. are both absorbed through the cell wall. two substances: water and combining very simple inorganic carbon dioxide.UNICELLULAR ORGANISMS of light 19 and by virtue of its chlorophyll. have divided to form small groups of cytoplasm.

ovoid in form and about 1/3. but separation usually occurs shortly after division.20 FUNDAMENTALS OF BIOLOGY cell in half. The nucleus The bud now enlarges and becomes pinched off from the parent cell. is cell visible in the living. called Saccharomyces cercvisiae. and a thin cell wall composed of but there Each oil many by small no chloroplast. Soon the chains break up. A bud outgrowth. Reproduction occurs arises as a small budding. It merely involves an approximately equal division of an individual into two new ones that become independent organisms. the common yeast plant is unicellular. cutting the increase in size. but is It has a nucleus. short chains of cells being formed. . colorless. cytoplasm. each Fission is the simplest cell taking up an independent existence. each arising from a single cell. The two new cells now either immediately separate. Yeast. the two nuclei. sometimes even and more. one of the commonest. a unicellular colorless plant. Reproduction occurs by budding. it may either separate at once or remain attached and produce another bud. usually appearing at one end of the cell. 6. 6). Yeast. method of reproduction. a modified form of fission. one going into the bud. Thus from a single individual a group of four cells may arise. the cell being entirely contains one or more large vacuolcs and The nucleus is very small and not globules. each cell carrying on a separate existence. or frequently remain together until each has again divided. FIG. X 1. being used in making bread. unstained condition. Like Protococcus.500. cellulose. divides to form two nuclei.000 inch in length (Fig. There are a number of kinds of yeasts. In this way short chains may be formed.

for Fermentation in yeasts is accomplished by the production of an enzyme It is most active in the absence of free oxygen and serves called zymase. Bacteria are unicellular plants lacking chlorophyll and 1 live. and small amounts of other substances.500. in the air. 7. Spirillum undulalum. absorb. absorbing it in solution through the cell wall. it is called a saprophyte. ethyl This process is Bacteria. -The bacteria are at once the smallest and the They are found under all simplest of all known organisms. fiarcina lutea. Yeast cells use as food only a very small proportion of the sugar that they self-sustaining. and in the bodies of other organisms. Lacking It unable to carry on photosynthesis. JD. A.UNICELLULAR ORGANISMS The yeast 21 plant lives in fruit juices and other sugar solutions. source of food. D FIG. Bacillus F. Streptococcus pyrogencs. as a means of releasing energy when the ordinary type of respiration cannot be carried on. Protococcus is said to be an independent plant because it has the power of making its own food and is thus chlorophyll. It simply lives on sugar that is already made. taking in its food by absorption. The rest is alcohol. E. in soil. Bacillus Spirillum cholerae. B. requiring an external Because it lives on dead organic matter. 1 broken down into carbon dioxide. called fermentation. *" of e X 1. Group common typhosuGf bacteria. resembles Protococcus in utilizing sugar as a source of nourishment but differs in that it cannot make its own food. . They are of tremendous importance to man. it is Yeast is a dependent plant. conditions where life may exist in fresh and salt water. subtilis. not only because of their relation to disease but because of their beneficial activities as well. C.

Amoeba. While all bacteria are active only in the presence of moisture and other favorable conditions. 1 Some organisms. and curved or spiral (spirillum) forms cells (Fig.125 of the latter. single colorless cell.22 FUNDAMENTALS OF BIOLOGY the most part. about 1/10. It is a existence is known only by the effects they protoo small to be seen with the most powerful microIn fact they even pass through the pores of a porcelain filter. being widely Its body consists of a distributed in stagnant water (Fig. while in others they remain together permanently Such groups of indito form chains. This is one of the simplest types of unicellular animals. whose still duce. the best- known one. so that. There are a number of species of Amoeba. finger-like lobes extending in all directions. while many only 1/50. In some forms the two cells separate after division. in the course of 24 hours. Such a rate of multiplication is by the exhaustion of the food supply or soon checked. 8). Some are non-motile.000 inch very rapidly. or irregular masses. a single bacterium may give rise to billions. many bacteria can pass into a resting stage. There is no organized nucleus. 7). by means bacteria are so simple that they might almost be A thin cell wall surrounds a mass of said to be structureless. plates. are smaller . placed end to end. and scopes. remaining inactive for a long time. while others of bear slender which they move protoplasmic threads called cilia. rodshaped (bacillus) forms. Their are of three general types: spherical (coccus) forms. commonly called Amoeba proteus. Bacteria reproduce entirely by fission. if these fail.00( spherical inch in diameter. or on other living organisms as parasites. which is approximately j/fe inch in diameter. to stretch across the head of an ordinary pin. either on dead organic matter as saprophytes. in both cases absorbing food directly through the cell wall as in the yeast plant. however. viduals arc called colonies. Bacteria present on dust particles in the air are in a dormant state and can resist drying and great extremes of temperature. cells of The homogeneous protoplasm. for this reason are called filterable viruses. but merely some scattered granules of chromatin. irregular in outline because of numerous temporary. 1 It would take 625 of the former and 3. Under favorable conditions cell division may occur as frequently as every 20 minutes. by the accumulation of poisonous waste products of metabolism. The rod-shaped types average are forms the of in length.

and a beautiful demonstration of the power of independent motion. active. being about J^oo inch in diameter. X 250. a very thin plasma membrane on the outside. endoplasm. which is clear. Amoeba proteus. the central granular mass as Corrfracfite vacuole Pseudopodium Nucleus Food vacuo/es Ectoplasm Plasma membrarh \ ^ ^ Endoplasm Fia. 8. the is When flowing of the cytoplasm is a very striking thing to observe. With the exception of a comparatively thin layer on the surface. cell. termed pseudopodia.UNI CELL ULA R ORGA NISMS 23 relatively large cell. when it is seen to consist of many coarse granules. tion. living condition The is nucleus is rather difficult to see in the cell but very distinct after the has been killed and stained. . but most As in a typical animal other species are considerably smaller. there is a nucleus surrounded by cytoplasm. When the continually changing because of the extension and withdrawal of these finger-like lobes (Fig. large clear contractile vacuole is present in the cell. the animal slowly moves from place to place. a simple unicellular animal. and at the same time other pseudopodia are retracted. It A The animal finger-like extensions of the cell. while a cell wall and chloro- plasts are lacking. As a result of this The behavior. 9). is the cytoplasm clear layer known is finely granular. are the means is by which locomotion is is accomplished. suddenly contracts and then slowly expands. this action taking place regularly at short intervals. The outer as ectoplasm. its outline one or more pseudopodia are thrust out in a given direccytoplasm is seen to flow into them.

") Digestive fluids pass into the food vacuole from the surrounding cytoplasm.ter is taken into the cell with the food particle. E. outlines of a moving individual sketched at intervals to show the formation and withdrawal of pseudopodia. it may come It then thrusts out pseudopodia. other smaller animals. Fia. for otherwise life cannot exist. of the amoeba consists of unicellular green plants. oxygen is absorbed from the surrounding medium in order that the process of respiration may go on. to be characteristic of all living matter. D.24 FUNDAMENTALS OF BIOWOY of the which has been mentioned as one matter. A. (After Calkins. A small amount of wr. flow together. unique features of living The food bacteria. B. C. and of dead vegetable and animal matter. which surround the food. the assimilation. and engulf it. The . forming a food vacuole. Ajs in one-celled plants and all other living cells. F. 9. The digested food material. being merely left behind as the animal moves on. Locomotion in Amoeba. The work of digestion now begins. As the animal slowly creeps along. increases the amount of protoplasm in Indigestible matter is expelled from the body at any part of its surface. by cell. and the food is gradually changed chemically in such a way that it can be absorbed and thus made part of the This is the process of assimilation that we have seen living cell. in contact with a food particle.

ucts to be formed. These are chiefly water. but. carbon dioxide. 10). tion of organic matter in the cell causes metabolic waste produrea. (From Skull. from heat. direction.") the it cell. or from food is near. eonstiicof the cytoplasm. beyond which the animal does not go. more protoplasm is built up by assimilation than is destroyed by respiration. 10. When an active amoeba in is touched with a solid object. showing the division of the nucleus. the cell dividing into two parts. and then reproduction The amoeba. Then these move apart. each of which takes up an independent existence (Fig. like most other unicellular organisms. irritating chemical. and as a result growth takes place. Fission in Amoeba. the animal usually moves in its These responses are manifestations of irritability. just as when a piece of coal is burned. and . strong light. Thus this structure plays a part in the work of excretion. A size limit is soon reached. First the nucleus divides to form two nuclei. If the amoeba gets plenty of food. Similarly it by moving moves away from an if reacts the opposite direction. occurs. reproduces by fission. and separation of the two new cells. It also serves as a means of getting rid of excess water taken into tioii FKJ. Urea is the cell directly through the plasma membrane. while some apparently flows into the contractile vacuole and is expelled by it through the plasma membrane into the surrounding water. "Heredity. however.UNICELLULAR ORGANISMS 25 oxygen combines with the protoplasm with the result that energy is The oxidaliberated. and a nitrogenous substance resulting from the decomMuch of this waste matter passes out of position of proteins.

as reproduction by age is reached. 11. layer around itself.26 FUNDAMENTALS OF BIOLOGY between them the cytoplasm gradually becomes constricted into two approximately equal parts that finally separate to form two new cells. A cell curious fact about one-celled organisms is that. Cilia Pellicle Contract!le vacuole Oret /groove Micronucleus Oral funnel Ana/ pore FIG. is a unicellular animal. . the animal passes into a quiescent state. Paramecium caudatum. although the may die through accident. In the presence of adverse conditions. as during a drought. becoming spherical and secreting a protective Such an individual is said to be encysted. It remains in this state until revived by the return of favorable conditions. natural death docs not occur. Each individual fission takes place before old potentially immortal.

Two nuclei ways are present: a large oval macronucleus and In cell structure groove. the send the new course is now is the unfavorable stimulus encountered. as when an obstacle or an irritating cilia substance is reverse their action and animal backward. but common in stagnant ponds and streams (Fig. Paramecium is more or less slipper shaped. The shaded areas represent currents of water drawn into the oral Paramecium differs in from Amoeba. commonest species. One of the caudal urn. protoplasmic threads called cilia. but if A encountered. but swims in a spiral path. by per- . fine. 1 to 4. several "Behavior of the Lower " C olumbia Organisms University Press. like it is 27 This is unicellular animal than the a more complex amoeba. 12. called the again trial and if error observed study. being rounded at the anterior (forward) end and pointed at the posterior (rear) The cell is entirely covered over end. method. 12). the cell has a definite and constant form. being about 3^25 inch long. at the same time rotating on its long axis from right to left (Fig. taken. may easily be living animals are available for FIG. but under special circumstances. 11). Thus the method of locomotion in Paramecium and Amoeba is very different. with very short. which beat against the water and rapidly propel the animal forward.UNICELLULAR ORGANISMS Paramecium. a single individual is just barely visible to the naked eye when seen against a black background. Diagram showing spiral path taken by a paramecium and rotation on its long axis from right to left. Owing to the presence of a firm outer membrane called the pellicle. . This type of behavior. (From Jennings. In fact. the same avoiding reaction is repeated until a clear path is found. successive positions. It is also a large form. but there is no cellulose cell wall. Ordinarily the animal swims with the rounded end forward. Paramecium does not take a straight course through the water.

The cell also at either end. This breaks from the end of the funnel and passes into the cytoplasm. and in the constriction of the cytoplasm. one located Usually these alternate in contracting at intervals about 10 to 20 seconds. Paramecium caudatum has one micronucleus. forming a food vacuole. but some and some have several. In the anterior half of the animal is a shallow depression called the oral groove. Fission in Paramrcium. These seem to collect liquid waste products. .28 a FUNDAMENTALS OF BIOLOGY small spherical micronuclcus.Extending radially into the cytoplasm from each contractile vacuole are a number of minute canals. 13. consisting of bacteria and smaller one-celled animals. ABC emptied into the vacuole and finally discharged to the outside. plasm termed the oral funnel. species micronuclei. showing successive stages in the division of the inacronudeus and niicromicleus. which are Fid. which extends diagonally backward from left to At its lower end is a funnel-like opening into the cytoright. The oral groove is lined with long cilia whose beating creates currents that sweep food particles. A number of food particles enter the cell in loose 1 a drop of water. have two . The contractile vacuoles also relieve the cell of excess water. of contains two contractile vacuoles. 1 These are in contact with each other and have a constant position near the center of the cell. into the lower end of the oral funnel.

Reproduction When Both the macronucleus and the micronucleus elongate and divide*. processes of digestion. Indigestible matter is expelled through a definite anal pore that opens into the oral groove. and extending inward is a deep oral groove also bordered by Since the animal is not free swimming.UNICELLULAR ORGANISMS 29 By a streaming movement of the cytoplasm. second contractile vaeuole arises in each and A daughter fission. by fission increase to the size of the original Vorticella. One cell has the original finally separates into two halves. in used locomotion. These are trichocysts. 14). occurs by fission (Fig. a definite spot on the surface of the body. they discharge into the water long hair-like threads that form a protective network and tend to keep away another animal attacking the paramecium. oral groove leads to an oral funnel where food particles (chiefly bacteria) accumulate and finally enter the cell in the form of a food vaeuole. A peculiar feature of Paramecium is the presence of numerous is specialized. a small micronucleus. the cilia are not cilia. cell. sac-like structures in the outer part of the cell (the ectoplasm) just beneath the pellicle. Paramecium there is called the anal pore. oral groove and oral funnel. respiration. the 1 The peculiar behavior known as conjugation is briefly described elsewhere (see p. and a long U-shaped macronucleus. where indigestible matter is expelled. . 229). a form related to Paramecium. but by their rapid beating create a vortex As in Paramecium. the into which currents of water are drawn. living in ponds and ditches. when each may again undergo Another common unicellular animal. The two individuals formed cell. globular Vorticella has one contractile vaeuole. and excretion are carried on in essentially the same way as in Amoeba. The body of Vorticella campanula is bell-shaped It is attached to an object in the water by means of a (Fig. Reproduction occurs by fission. the food vacuoles The following a definite course. assimilation. long contractile stalk. Around the upper margin is a row of cilia. each forming two daughter At the same time the cytoplasm constricts transversely nuclei. but in pass to other parts of the cell. 1 13). but showing certain interesting differences as a result of its peculiar mode of life. This situated directly behind the lower end of the oral funnel. while the other half develops these structures anew. is Vorticella. stimulated.

is oral funnel. but some flagellates are amoeboid. often represented mic stagnant 15). is At the base a red pigment thought to be senorganism tends to swim toward the bestis which sitive to light. no food is ordinarily taken in through it. this organism has a rather definite form. micronucleue. are in One such pools group the flagellates. Vorticella campanula. Like Paramecium and Vorticella. but. that lashes back and forth and pulls the organism through the water. encystment may body dividing Euglena. Instead of ingesting solid food particles as unicellular animals do. there are some low forms of life with mixed affinities. o. 14. swims by means whip-like of long. conditions. food vacuole. A a opens into a permanent cavity single nucleus is present in the posterior part of the water. gradually tapering behind. wall is A cellulose is not its present. termed the reservoir. mac.30 FUNDAMENTALS OF BIOLOGY In the presence of unfavorable longitudinally. as the Fio. Euglena makes its own food. At the anterior end of the body is a small oral funnel that leads to the reservoir. putting out slender pseudopodia. Although it is possible to assign most organisms to either the plant or the animal kingdom. occur. for within the . strangely enough. macronucleus. contractile fv. . mic. cv. cilia. stalk. cilium (usually termed a flagellum). owing to the presence of a thin elastic pellicle. c. an attached unicellular animal. anteriorly. illuminated although light. vacuole. avoiding direct sunNear the pigment spot contractile vacuole that part of the cell. X 600. combining characters of both plants and animals. of the flagellum spot. Euglena It blunt at forward a single attached end. and ditches by a form known as Euglena mac viridis (Fig.

particles of food through the oral funnel. ericystment in the presence of unfavorable conditions a common occurrence. Other flagellates are saprophytic. Thus its nutrition is distinctly Reproduction occurs by fission. . they become colorless and saprophytic. Some species'of Euglena carry on photosynthesis but if kept in darkness and supplied with organic material.) has chloroplasts and carries on photoSome forms are synthesis. -F/agellum -Oral funnel Reservoir- -figment spot -Confracf/le vacuole * Chhroplash -Nucleus FIG. the other longitudinally. certain other flagellates are colorless. (After Doflein. forming a new one. Euglena viridis. absorbing liquid Some take in solid organic matter as the yeast plant does. is As in Amoeba and many other unicellular organisms.UNICELLULAR ORGANISMS 31 cell are a number of small chloroplasts that enable the organism to carry on photosynthesis. while others engulf food an amoeba. which causes Although Euglena viridis sickness. African sleeping parasitic. the cell dividing plant-like. One half keeps the old flagellum. an organism showing both plant and animal characters. like in the light. 15. such as Trypanosoma.

Thus the relation of flagellates to the two great letter organic kingdoms may be likened to that of the stern of the Y to its two branches. a simple chain-like colony with a differentiated cell marking the place where the colony will break. D.) FKJ. by emphasizing the acquisition of food from plants. emphasizing photosynthesis. the cells imbedded in a mucilaginous matrix. a spherical. flexible tissues. representing a common ancestral slock from which later both plants and animals may have arisen. Anabaena. suggests that the first forms of life may have been similarly undiffercntiated. on the other hand. cells in close Some contact. It has been seen that cell division in uniorganisms results in the production of two new indi- . (B. and has arisen in consequence of that primary cellular Colony Formation. and motility. C. features of plants and animals is related to the basic factor of nutrition difference. Pandorina. fixity of position. X 750. intermediate between plants and animals. a simple colony. the liberation of Each of the distinguishing energy. Pediastrum. Plants became differentiated by simple plant colonies. storage of energy. 16-celled. a plate-like colony with X 750. X 1. C and D. Glococapsa.000. B. motile colony enclosed in a mucilaginous sheath. 16.32 FUNDAMENTALS OF BIOLOGY The occurrence of such organisms as flagellates. A and B are green algae. after Pringsheim. and Animals arose. rigid tissues. A. blue-green algae.

the cells produced by fission show a As a rule. may be arranged in chains. cells forming a colony spheres. The ments. but uncommon among one-celled animals.) depending on whether the cells separate following division or remain permanently together. Carried further. this tendency would result in the formation of A colony is an aggregation of individuals. A. 17. each maincolonies. masses. tendency to remain in contact for a while before separating. B FIG. after Ktdn\ B.UNICELLULAR ORGANISMS viduals. being confined Chiefly to forms related to Vorticella (Figs. but in such forms as Protococcus and yeast. 33 these immediately separate. a protozoan. Potcriodendron. 16 and 17). by moans (A. a flagellate. irregular otherwise. or fila- plates. Tree-like colonies of unicellular organisms held together fi. The . Thus unicellular organisms may be either solitary or colonial. after Kent. of stalks. Epistylis. taining itself and having little or no dependence upon the others. Colony formation is rather prevalent among unicellular plants and flagellates.

on the one hand. and multicellular organisms intergrade is of great significance. It is only under special circumstances that a cell may become separated from the organization and carry on an independent existence. some cases certain cells. It is apparent that no sharp distinction exists between the more highly organized colonies of unicellular individuals. all the cells in the colony are alike. ordinarily this is not possible. In more highly developed colonies the cells are in contact with one another. and so there is organic union between them but no physiological dependence. thus resulting in a and animals are multicellular. become differentiated. In fact. In the evolution of the early forms of life. Another feature of multicellular organisms is the vidual of multicellular organism. the tendency for cells to any line of The fact that unicellular remain together after division led to the development of colonies. or by some other mechanical means. Here each cell is but a small part of the individual.34 FUNDAMENTALS OF BIOLOGY simplest colonies consist of cells held together very loosely by a mucilaginous matrix. . All the higher plants fact that the cells of the their functions body cooperate in the performance of and are more or less dependent on one another. with specialized functions. others of many thouAs a rule. as any cell may become detached from the colony and continue to maintain itself. by stalks. and simple multicellular organisms on the other. Some colonies consist of only a few cells. the indicells closely associated in innumerable consisting structure and function. demarcation drawn between them would be arbitrary. but in sands. while a closer association of the cells in the colony may have led to a dependence of the cells on one another. indicating that the latter may have been derived from the former through the formation of colonies.

forest. The plant kingdom is divided into four major groups.CHAPTER IV THE LOWER PLANT GROUPS In discussing organisms of greater structural complexity than the unicellular forms considered in the last chapter. and garden. most of which are structurally complex. and classified by botanists. THALLOPHYTES The of simple plants thallophytes constitute a large and diverse assemblage forming the lowest division of the plant kingdom. Over 300. and each succeeding group representing a higher condition of structural organization than those below it. simpler than animals.000 of one or of many cells body may consist most (although thallophytes are plant The multicellular) but in the latter case is nearly always a thallus. thallophytes comprise two subordinate groups: the algae and the fungi. species. forms. ALGAE respect to their nutrition. This chapter will be devoted to a consideration of the three lower plant groups. roots. it will be more advantageous to treat of plants and animals separately than to attempt any other plan.000. They are the simplest phytes. all of With 35 . in general. that is. the latter 70.. viz. The former number 20.000 different species of plants have been* named..000 species. while the two that follow will deal with the highest group. and leaves. includes not only the various kinds of familiar plants of field. but also the many simpler ones largely unknown to most people. a plant body with little or no differentiation into distinct vegetative organs. Plants will be studied first because they are. cell They number about 90. It is chiefly to The this feature that the thallophytes owe their simplicity. collectively constituting the plant kingdom. This vast assemblage of described. algae are independent thallothem having chlorophyll. the members of each having certain basic features in common. stems.

16. Many other unicellular forms occur. A few of the latter are shown in Fig. Cvlenchaete. forms. a typical one-celled alga. X 250. A. are microscopic. the . Many size. a simple unbranched filament with a differentiated basal cell that serves as a holdfast. Among the simpler multicellular algae.FUNDAMENTALS OF BIOLOGY of all green plants and are thought to be also the oldest. kelps and other seaweeds. X 150. but some of the seaweeds reach a large Vegetative Body. Drapcirnaldia. a branching filament showing differentiation in size of cells. It. They include the pond scums. others colonial. (\ Ulothrijc. Protococcus. Some simple multieellular green algae. both fresh and salt. the cells in a plate-like airangement. the small bodies on the chloroplast (pyrenoids) are centers of starch formation. X 350. 18. has already been discussed. In B and C the peripheral band-like chloroplast obscures the central nucleus. some of which are solitary. and many less familiar Fio. Nearly all the algae live in water.

the algae are also relatively simple plants. the prevailing method of reproduction is by fission obviously the most primitive method . In many cases all the cells of the body are essentially alike in form and structure. but often there is a tendency for some of them to become different. Structural differentiation becomes marked in the higher plants. 19. differentiation in size. A. while some algae consist of a thin plate of cells (Fig. D. formation of spores inside the vegetative cells. For example. hand-like chloroplast with a number of starch-forming bodies (pyrenoids) B. eventually leading to the formation of tissues and organs. and leaves of the higher plants. as in vegetative Among the features. 23C). unicellular forms. 18C). but which structurally are much simpler (Fig. In many of the brandling forms the cells A Fio. show a Asexual Reproduction. such as Protococcus. X 700.- " 11 B C . -Reproduction in Ulothrix. 18#). a simple green alga.THE LOWER PLANT GROUPS body commonly lias 37 the form of a simple filament (a row of cylindrical cells attached end to end) or a branching filament. formation and escape of gametes. (\ free-swimming spores which have escaped. portion of Kaeh cell contains a central nucleus and a peripheral vegetative filament. In reproduction. some of which are pairing. one or more cells may be modified to serve as a means of attachment to an object in the water (Fig. stems. The most highly differentiated algae are marine forms with a body consisting of parts that bear a superficial resemblance to the roots. those of the branches being smaller than those of the main filament (Fig. 18).

. become and then The pairing The cell that . behavior being called germination. These escape from the plant cell wall) and by means of A fact of special interest in con- nection with these that many of swimming spores is them resemble flagellates. Sexual Reproduction. they are incapable of directly a new plant. these resemble swimming spores in appearane'e but are usually smaller (Fig. spores are 1 produced is in specialized structures called sporangia. Although gametes arise algae is 1 in the forming same way that spores elo. is by the production of swimming cell spores. but it is only among unicellular division reproduction. 20. FUNDAMENTALS OF BIOLOGY All cells cell divide. not only in being naked and motile. 20. pigment spot (see pp. Tn many of the simpler algae.1).38 possible. may give rise to a new X 400.Sexual reproduction among the lower a relatively simple process. 1979). A 1 spore may be defined as a having he capacity of directly developing into a new plant. whose sole function spore production (Fig. each spore. as naked cells (lacking a swim through the water cilia. Among organisms that multicellular organisms it results in growth. and reproduction takes place by the separation of a cell or a group of cells from the parent. algM. they come The essential feature of sexual reprotogether in pairs and fuse. After coming to rest. Instead of germinating. the new individual developing from this fragment. plant by cell division. but in having a contractile vacuole and a B F[(j. 19 and 22). this :i single osrjipod sporo. Sporangium Also (A} and KnmptuMgium (#) of a brown Kcfociirpus. the contents of any one 1 of the vegetative cells may give rise to one or more uninucleate swimming spores (Figs. 30 and 31). detached cells called gametes are formed. all duction in cells organisms one. Naked. In many of the algae. their nuclei unite. A common method by which this process operates in the results in algae. Sporangia are characteristic of all plants above the thallophyte level. is the fusion of two gametes.

later either giving rise to a new plant directly. X 250. C. a vegetative roll showing a central nucleus and a spiral ribbon-like rhloroplast with many starch-forming bodies (pyreiioids). . FIG. In some algae two filaments. D. si ages in sexual reproduction. B. forming tubes from one filament to the other (Fig. gives rise to a single large gamete. itself algae usually secretes a heavy wall around and goes into a dormant condition. X 500. BCD ciliated or escape into the water. . or to four spores.THE LOWER PLANT CROUPS arises 39 In the from the union it of two gametes is called a zygote. forming heavy-walled zygotes. 21). lying parallel to each other. which may be active or passive. 2 1 --Spirogyra a common green alga. but does not become This . put out lateral projections that come in contact. each of which in turn produces a new plant (Fig. Through these tubes the cell contents of the one filament pass to fuse with those of the Here each vegetative cell other. A. 22F).

after Ilirn.) BCD E only a relatively few algae are both of the fusing gametes and behavior.'. 22. Fid. F. (.walled zygote resulting from the fusion of sperm and egg. a green alga. female filament with a large egg-producing cell and a vegetative cell. E. after Juranyi. although not a typical alga in many respects. A and B.1 and B. F. the entire contents of a vegetative cell escaping as a single swimming spore. The one remains small and active. (\ malo filament with two groups of sperm-producing cells sepaiated by a vegetative cell. while the Ill alike in size other becomes large and passive. and numerous starch grains.40 FUNDAMENTALS OF BIOLOGY peculiar method of sexual reproduction is mentioned because it takes place in tipirogyra. In most plants gametes are of two different kinds. also a single escaped sperm. the egg contains a nucleus. starch-forming bodies (pyrenoids). D. Keproduction in Oedogonium. group of four swimming spores produced by the zygote. one of the commonest of fresh-water it is algae and a form widely studied in elementary courses. nearly always being retained . a heavy. X 500.

These are the lowest of the algae. the cells contain a definite nucleus and one or more distinct chloroplasts. The larger one is known as the egg or female gamete. or by gametes. water forms in which chlorophyll is the only pigment present. this act being called fertilization. in gametes are produced special cells known as gamctangia or sexual organs (Fig. 2()#). tribution. the pigments being merely diffused throughout the cytoplasm. called sperms or male gametes. except in one group. As in all the higher algae. the algae are classified. in addition to the chloro- They are all solitary or grouped unicellular plants. occurring along nearly all seacoasts but reaching their . Blue-green Algae. The sperm swims to the egg and fuses with it. whether differentiated into sperms and eggs or not. These algae are almost all marine in dis3. 5. Like sporangia. Their color is due to the presence of a blue pigment phyll. and 22). its motility being sacrificed for an increased nutritive capacity. In the central part of the cell is a very primitive nucleus consisting only of a mass of chromatin granules not sur- rounded by a nuclear membrane. 1. are ciliated and escape into the water. Repro- duction occurs exclusively by fission. these are formed cells just but. Chloroplasts are not present. 18. but unicellular forms occur and may be either solitary or colonial (Figs. arise directly from ordinary vegetative In maify of the algae. The gametes of the lower algae. 16(7 and D). The four groups of algae. by spores. Green Algae. In cell structure this group is the most primitive of all green plants. of a The often zygote resulting from the union sperm and an egg is called the fertilized egg. 19. 22(7 and Z>). are given below. Most of them are multicellular. Reproduction may occur by fission. with their distinguishing characters. All the algae contain chlorophyll. some other pigment is present that more or less obscures the green color. living in both fresh and salt water and on moist soil. The large size of the egg is due to the accumula- tion of reserve food in the cytoplasm. 21. The smaller gametes. Groups of Algae. 16/1 and /?. and the cells may be either to form colonies (Fig. and it is chiefly on this basis that especially for the production of reproductive cells. Brown Algae. as the spores do. however.THE LOWER PLANT GROUPS within the cell 41 from which it arises (Fig. -The green algae are predominantly fresh2.

X M. plate-like. a plate-like red alga (Grinnellia) a delicately branching red alga (Gelidium). in addition to the chlorophyll. D. a small kelp (Eisenia). filamentous.42 FUNDAMENTALS OF BIOLOGY Their cells contain a greatest development in cooler waters. * % brown algae are unicellular. None of the brown pigment Some common seaweeds. X }*>. a rockweed (Hesperophycus). X Fia. A. or may and the multicellular body may be reach massive proportions and . # . X H. 23.

it is in this group that the filamentous type of plant body reaches tion takes place its greatest development. (From Sinnott. Red Algae. Reproducand by spores gametes. Some growing along our Pacific Coast reach a length of 100 Reproduction occurs by means of both spores and gametes.") pigment is present in the cells. sporangiophoro. />. or forming plates (Fig. No unicellular forms are known. C. 4. ribbon-like. 24. 20. being filamentous. 234 and B). "Botany. Like the brown algae. The filamentous red algae are highly branched and very delicate. sexual reproduction being very complex. form (Figs. horizontal branch. A. B. and the multicellular bodies are not large but very diverse in shape. sporangium. All of them lack chloroand live either on dead organic matter as saprophytes. a red Fio. haustorium. Principles and Problems.THE LOWER PLANT GROUPS become differentiated in 43 of the kelps to 150 feet. Mycelium of bread mold (Khizojms) giving riso to sporangia. the red algae are almost exclusively marine but as a rule live in deeper and warmer waters than do the browns. In addition to chlorophyll. In fact. FUNGI phyll Fungi are dependent thallophytes. 23C and D). .

molds. and mushrooms. in Rhizopus. yeasts. food is absorbed (Fig. the common called a mycelium. white mold that grows on moist stale bread. B. sporangium of a water mold (Saprolegnia) with swimming spores. but in nourishment directly from living In the group of fungi to which the bread mold belongs. such as the water molds aquatic forms that live on dead fishes and insects swimming spores are produced. smuts. rusts. In parasitic fungi the mycelium may of the organism attacked.44 FUNDAMENTALS OF BIOLOGY Familiar kinds of fungi are blights. sporangium of bread mold (Rhizopus) with aerial spores. In some of the lower fungi having a mycelium. 25. the mycelium does not have cross walls dividing it up into individual cells. but in most fungi cross walls are present. The bacteria and other unicellular fungi reproduce by fission. 25). but practically The all bacteria and yeasts are unicellular other fungi have a thread-like. grow on or beneath the surface either case the fungus absorbs cells (Fig. forms. or on living organisms as parasites. mildews. the mycelium consists of a fluffy mass of delicate threads that send short root-like branches (haustoria) into the bread and through which body FIG. bacteria. 24). branching For example. Aerial spores differ from swimming spores in lacking cilia and in having . Reproduction. Spore production in two common fungi. Vegetative Body. A. 26). but in the majority of fungi the spores are aerial (Fig.

27). but in both oases the mycelium absorbs ^^^r B nourishment directly from living cells. /?. a mushroom (Amanita) that lives in soil. Two parasitic fungi. and produce innumerable aorial spores. conspicuous fleshy bodies that produce Sexual reproduction among . X J. a smut (Ustilago) attacking an ear of corn. some of the grains of which The mildew is an exterare greatly enlarged and filled with black spores.THE LOWER PLANT GROUPS a cell 45 wall. a mildew (Erysiphe) growing on a bean leaf /^. 26. the smut an internal one. Two saprophytic fungi. B. fungi the spores are borne in In many of the higher forms A X FIG. the mycelium gives rise to innumerable aerial spores (Fig.In. both cases these fleshy bodies arise from a colorless mycelium living in the decaying organic matter. ^PT B A. 27. X M. a cup fungus (Peziza) growing on rotting wood. A. In practically all sporangia or in similar structures. A 'FiG. X J^. nal parasite.

46 FUNDAMENTALS OF BIOLOGY of the fungi is many an obscure process and is frequently not present. X 2. while the bryophytes. flat thallus with upright branches bearing sexual organs. 28. male and female plants. rather inconspicuous. X 1. being closely related on the one hand to the green algae and on the other hand to the higher plants. It is noteworthy that the simplest plants (the algae) are aquatic. of a liverwort (Marchantia). but . Bryophytes are small. A and B. BRYOPHYTES The second mosses. and in similar situations. Two common bryophytes. C. The long-stalked sporophyte is attached to the upper end of the leafy gametophyte. The gametophyte consists of a respectively. a moss (Funaria). comprising the liverworts and species. numbers about 20. considerable scientific interest in that they are a transitional group. green plants that live mostly on moist They are of soil. FIG. tree trunks.000 great plant group. rocks.

while in the mosses they arise in clusters at the apex In some mosses both kinds of sexual of the stem (Fig. are mostly unicellular and simple. The latter swim toward the archegonia and enter . The sexual organs of tliallophytes. which are both anchoring and absorptive in function. In most of the liverworts and in all of the mosses. neck of the archegonium opens. forming a passageway to the egg. the female organs archegonia. each having a slender neck and a lower bulbous portion consisting of one or more layers of sterile cells When mature. In the bryophytes. made up largely of simple uniform cells. while the cells of the leaf generally form a single layer. ellipsoidal organs consisting of a layer of sterile cells surrounding a large number of small sperm- The male organs producing cells cells (Fig. Sexual Organs. in others. Some of the liverworts have a vegetative body consisting of a flat. cells or may This may consist of only a few layers of simple be thicker and show some internal differentiation. the enclosing a large non-motile egg (Fig. It is apparent that an erect leafy shoot permits of a greater exposure of green tissue to the light than does a flat thallus and thus favors photosynthesis. contains no woody tissue. and in none of the bryophytes are roots present. 29 B). whether thalloid or leafy. 29A). grow on land but mostly in wet places close to the ground. 28(7). where present. filamentous hairs called rhizoids. The antheridia are stalked. sends into the soil numerous slender. gametes are borne in complex sexual organs that arise on the vegetative body described above. The sperms are minute curved bearing a pair of long cilia (Fig. The stems and leaves of bryophytes are much simpler than those of the higher plants.THE LOWER PLANT GROUPS 47 slightly more advanced structurally. bryophytes arc called anthcridia. In the presence of moisture the antheridia break open to liberate the sperms. organs are present in the same cluster. but those of all bryophytes are multicellular and complex. plate-like or ribbon-like thallus (Fig. The archegonia are flask-shaped organs. 28A and B). 29(7). exhibiting a of characteristic structure. In some of the liverworts the sexual organs are borne on special upright branches of the thallus (Fig. they occur on separate plants. Vegetative Body. 28 A and B). The vegetative body. the vegetative body is not a flat thallus but is made up of simple stems and leaves (Fig. The stem. 29D).

longitudinal section of an antheridium. The latter comes from the zygote and in turn produces spores. Life history of a moss (mainly Polytrichum). //. FUNDAMENTALS OF BIOLOGY Fertilization is effected it when a single sperm pene- trates the egg and fuses with to form a zygote. viz. B. X 1. longitudinal section of an archegonium. however. A. it is called a gametophyte. 29. enlarged view of several spores. X 150. X 150. a protonema bearing a bud from which an erect leafy shoot will arise. A different kind Fio. . X 75. enlarged view of a sperm. D. E. F. of individual is involved in the life history of the bryophytes. a capsule from which the outer covering has been removed. Because the green vegetative body produces sexual organs in which gametes arise. a sporophyte. female plant with a mature sporophyte arising from its upper end.48 the necks. X 1. (?. C.. male plant bearing a cluster of antheridia at its upper end.

In the mosses. obtaining its nourishment directly from the gametophyte. 29ff). being very different from the simple sporangia found When the spore of a liverwort germinates. SPORANGIUM AOTHERIDIUM SPERM SPOROPHYTE ZYGOTE FIG^. 30. archegonium enlarges. among the algae. 49 pass into a The zygote does not escape and resting condition. the cavity of the constituting the embryo. The embryo continues to grow. the spore produces a green. 28C and 29S). By repeated cell division. Diagram illustrating the general scheme of alternation of generations in a moss or a fern. however. each of which may then give rise to an erect leafy shoot that later bears the sexual organs.THE LOWER PLANT GROUPS Sporophyte. and an upper spore-bearing The foot is an absorbing and capsule (Figs. anchoring organ. as in inside the archegonium most of the algae. branching filament called the protonema (Fig. and in nearly all bryophytes finally becomes differentiated into three regions: a lower foot. The life history of a liverwort or a moss introduces the phenomenon of alternation of generations. The seta often becomes long. . but develops at once where it was formed. Within the capsule are formed numerous spores that when ripe are disseminated by the air. a middle stalk or seta. it gives rise directly to the green vegetative body that later produces the sexual organs. Upon this buds appear. The capsule is really a sporangium but is multicellular and often highly specialized. carrying the capsule up into the air. Alternation of Generations. the zygote gives rise to a mass of undifferentiated cells As it develops.

surface. while in C the leaves are large and divided. stems. the gamotophyte the sexual one. is not because it derives this all or most of its nourishment from the gametophyte. X %. A definite alternation of generations is in all plants above the thallophyte level. An gametophyte important feature of the bryophytes is the fact that the is always an independent plant. In A and B the leaves are small and the sporangia borne in terminal cones. thoroughly established but notable changes . The former always arises from the zygote. C. this in of individuals alternate every life cycle. and leaves of the two higher plant groups. but the sporophyte FIG. one producing spores (sporophyte) and the other gametes (gamctophyte) Figure 30 represents the general scheme of alternation of generations. a horsetail (Efjuisetum). conspicuous. stems. a lycopod (Lycopodiu m) X Representative ptcridophytcs. A. reason the gametophyte is relatively It should be kept in mind that the rhizoids. Briefly stated. For and leaves of bryophytes belong to the gametophyte generation and have none of the complex structural features exhibited by the sporophytic roots. 31. 7?. X ^2'. the latter from a spore. a fern (Poly podium). and the sporangia borne in localizing groups on their lower . The sporophyte represents the asexual generation.50 FUNDAAfENTALS OF BIOLOGY means that two kinds .

PTERIDOPHYTES The group ferns and their relatives constitute a comparatively small of plants today. but frequently is not well defined. 32. 16. of a cross section of a fern stem (Ptcris aquilina).001) FIG.THE LOWER PLANT GROUPS 51 occur in the relative importance of the gametophytc and sporophyte in the life history. the " equisetums or horsetails. There are three modern groups of much more numerpfmdophylc*: the (often inappropriately called "club mosses"). but in the lycopods and horsetails the leaves are small. being represented by only about 6." and the ferns. and closely crowded on the . Fern leaves are typically large and divided into many small leaflets. as will be seen later. but in past geologic times they were ous. Photomicrograph X of The vascular system is composed xylem surrounded by phloem. An ordinary fern plant consists of an underground stem (rhizome) that sends out roots into the soil and leaves into the air. of numerous strands. each consisting species. lycopods Vegetative Body. the gametophyte and sporophyte being inde- pendent of each other and often similar in appearance. undivided. Alternation of generations occurs in many of the thallophytes. By far the largest group is the ferns.

but in some ferns no indtisium Each sporangium is a stalked organ consisting of a present. the archegonia occurring near the notch. 32). heart-shaped thallus that reaches a diameter of about. The vegetative body described above is a it to to the leaves. constituting the annulus. Spore Production. spores. are present in practically all pteridophytes. Sexual organs are also borne on the lower side. structure (Fig. which are relatively large. Xylem (wood). permitting grow up into the air. 34). two lower plant groups. much more complex structurally than those of bryophytes. cross section of a fern stem reveals the presence of a welldeveloped vascular or conducting system (Fig. When a fern spore falls upon moist earth. The sporangia of ferns commonly arise on the underside of the ordinary leaves in small groups called sori (Fig. 35). the antheridia being scattered among the rhizoids. A sorus is usually covered by a membrane known as the indusium. while which are entirely absent in the roots. 33). coiled. From its lower surface numerous rhizoids are put out into the soil. In all pteridophytes these are borne in complex sporangia. when mature. extends vertically about two-thirds of the way around the spois rangium. are discharged from the antheridia aad . but is characteristic of all the higher plants.}/ inch and closely resembles a simple liverwort (Fig. It gives strength to the plant. sporophyte because it produces spores. and multiciliate. beginning in the roots and extending into the leaves. made up largely of greatly elongated cells with thick cell walls but no protoplasmic contents. it is concerned with the discharge of the spores. green. layer of sterile cells enclosing a number of thick-walled aerial A ring of thick-walled cells. It consists mainly of two kinds of is tissues: xylem and phloem. A vascular system is not present in the thallophytes and bryophytes. 31). simpler in the ferns than in the mosses but have the same general The sperms. it produces a flat. from the leaves to other parts of the plant. 31/1 and B). This is A continuous throughout the plant body. Both kinds of sexual organs are Gametophyte.52 FUNDAMENTALS OF BIOLOGY stem The stems and leaves of pteridophytes are (Fig. and also conducts water from the roots Phloem consists of elongated cells with thin cell Its function is to transport food walls and with protoplasm. In the lycopods and horsetails the sporangia are borne in terminal cones (Fig.

portion of the underside of a leaflet showing the sori. and the zygote germinates at once. Since the thallus is 53 in contact with moist earth. enters the archegonium and fuses with the egg. . Spore-bearing structures of a fern (Poly podium californicum) the sporophyte. As in the mosses. A. the zygote does not escape but produces an embryo that develops for a while inside . sori arc not covered by an indusium. The sperm FIG. X 150.THE LOWER PLANT GROUPS swim toward the archegonia. 33. B. X %. a single sporangium with its annulus and several spores. In this fern the r. fertilization presents no difficulties. X 4. consisting of a rhizome bearing roots and large divided leaves.

35. longitudinal section of an aiitheridium containing a number of sperms. showing rhizoids and numerous antheridia. longitudinal section of an archegonium containing a single large egg. and three archegonia near the notch. X 350. more highly magnified. (0. B. Sexual organs of a fern. A. C.54 FUNDAMENTALS OF BIOLOGY FIG. X 500. X 35. View of the lower surface of a fern gametophyte. FIG. 34.) . after Yamanouchi. a single sperm.

THE LOWER PLANT GROUPS the archegonium. being leafless and entirely or largely dependent upon it for nourishment. soon forming a stem. but in some cases two kinds of spores are present. 30. the lattejj . The young sporophyte remains attached to the gametophyte for a while. FIG. the sporo- phyte attains independence and becomes the dominant generaThe fern gametophyte is consequently a greatly reduced tion. the sporophyte. and leaf. but soon becomes an independent plant (Fig. X 6. on the other hand. similarly produce small gametophytes that bear sexual organs Likewise the developof the same general type as those of ferns. The former condition is known as homospory. by It is and a fern developing highly differentiated vegetative organs. root. Young fern sporophyto attached to the gamotophyte. In most of the pteridophytes the spores are small and all alike. gametophyte ment of the embryo arid its relation to the gametophyte are essentially similar to the condition in ferns. apparent that the conspicuous difference between a moss is the presence of an independent leafy sporophyte in In the bryophytcs the gametophyte carries on all the latter. structure concerned with the production of gametes and not primarily with the manufacture of food. 36). Then the In the lycopods and horsetails the spores dies. absorbing nourishment from it. the latter being mainly a function of the sporophyte. or nearly all the work of photosynthesis. 55 The embryo grows rapidly. in all cases. In the pteridophytes. Heterospory.

several microspores and a megaspore drawn to the same scale are also shown. male gametophytes bearing only antheridia. X 25. which are sexually differentiated. 37). X 25. X 2. The microspores produce their sporangia and fall to the ground. C\ cone scale bearing a megasporangiuin containing four large spores. female gametophytes bearing only archegonia. showing terminal cones. of Selaginella willdenomi. the cone produces two kinds of sporangia. Both kinds of gametophytes are greatly reduced. the spores are shed from (Fig. a heterosporous pteriaophyte. one of the lycopods. . cone scale bearing a microsporangium containing many small spores. portion of sporophyto SeldQindla. C A. and a great reduction in the gametophyte generation. B. FUNDAMENTALS OF BIOLOGY Thus in Selaginella. Thus heterospory introduces into the life history not only two kinds of spores but also two kinds of gametophytes. the megaspores. B FIG. 37. the other giving rise to four large megaspores As in other pteridophytes. developing within the spore coat and remaining colorless. one kind containing many small microspores.56 as heterospory.

firs. angiosperms are a younger group than gymnosperms arid are much more numerous today. The name gymnosperm means "naked seed/ in reference to the fact that their seeds are borne freely exposed on the face of the cone scales (Fig.CHAPTER V VEGETATIVE ORGANS OF SEED PLANTS The spermatophytes of the plant is constitute the fourth kingdom. Like the pteridophytes they have an independent sporophyte with a woody conducting system and are also highly specialized in regard to the structure of their vegetative organs. The spermatophytes not only surpass the other groups in structural complexity. there being about 190. 57 . are an ancient group with a long geologic Like the pteridophytes. Historically. The cycads are tropical plants with large : fern-like leaves. but are the most numerous. They are often referred to as "flowering plants. with small needle-like or scale-like leaves. borne in a woody cone composed of scales (Fig. include the familiar pines. Nearly all the common plants of everyday experience are angiosperms. The former." as the presence of flowers is one of their outstanding features (Fig. redwoods. cedars. angio7 ' sperms are the dominant group of modern plants. hemThe seeds of gymnosperms arc locks. In fact. 67#). 67 A). while the conifers. Their name means "seed and highest division plants/' and appropriate because they are the only plants that bear seeds. There are two subordinate groups of spermatophytes the gymnosperms and the angio sperms. etc. 38B). 38A). The name angiosperm means "enclosed seed/' signifying that the seeds are produced in a closed vessel (ovary) that ripens to form a fruit (Fig. they were more numerous history. Gymnosperms are represented today chiefly by the cycads and the conifers. during past ages than they are now.000 species. nearly all of which are trees. numbering only about 500 living species. spruces.

The welfare of the plant demands that these be maintained at all times. A. all tive turally. functions are concerned with the nutrition of the plant body. a fruit which develops from one of the floral organs. collectively constituting its root system. of three distinct kinds of organs: roots. especialty dissolved mineral salts. a gymnosperm. tion. It also plant obtains its entire water supply through its roots. the vegelafeatures for the next chapter. dies. within. X ?2l ^ white trillium (Trillium In the pine two "naked" seeds are borne grandijlorum) an angiosporm. In the trillium many "enclosed" seeds are borne at the base of each cone scale. and leaves. we shall consider first thenvegetative organs. 38. providing for its growth and continued existence. body is made up stems. it soon wilts and eventually The primarily because its source of water has been cut off. mature seed-beat ing COUP and leafy branch of white pine (Pinus strobus).58 FUNDAMENTALS OF BIOLOGY In dealing with the seed plants. in intimate relations with the soil. . The root system performs two primary functions: anchorage and absorp- When a plant is uprooted. which are highly differentiated strucThey are called vegetative organs because their chief of B FIG. Spermatophytes. reserving a discussion of their reproductive As in pteridophytes. THE ROOT The place it roots of a plant. absorbs from the soil other substances necessary to its welfare. X /&. .

a root system grows. 39. elongation. the as it is only these delicate rootlets work of which carry on absorption. Root tip of a grass seed- examining with a microscope the young rootlets put out by grown in water. a. giving rise to many smaller branches. are many root systems of inter- mediate character. sprouting seeds (Fig. It soil in all should be noted that there the way in is no regularity to which branch roots the smaller ones springing from the larger ones without arise. Some root systems consist main root. which grows straight downward into the soil. its absorbing region moves farther and farther out into the soil.VEGETATIVE ORGANS OF SEED PLANTS Organization of Root Systems. Structure of Root Tip. that the larger roots give rise to smaller and smaller branches. and so these " " have no great signifitypes The important point is cance. called the taproot. pushing their way out into the soil. Fitting over the end of the root tip is a thimble-shaped group of cells forming the rootcap. a structure that serves as a buffer to the rootlet as it pushes through the behind. all of which are slender and equally prominent. region of d. c. . but the root system consists of a cluster of highly branched fibrous roots. protecting the delicate cells that lie immediately Just back of the rootcap is the embryonic region. rootcap. region of maturation. Furthermore. predetermined o r d e r. ling fr. Between these two extremes. X 35. The structure of a root tip can be studied to good advantage by a FIG. embryonic region. Another fact of importance is that the tips of all much roots are constantly elongating. In of one other plants one large root does not dominate the others. of soil. 39). however. and so. which penetrate the directions.

60 FUNDAMKNTALH OF JUOLOGY very limited extent. the cytoplasm no longer appears dense (Fig. seedling of scarlet runner bean with equidistant dots placed along the primary root. . Behind the region of elongation is the more extensive region of maturation easily recognized externally by the presence of . which are slender outgrowths from the outerIt is here that the cells of the cells (epidermis). consisting of a small group of undifferEach of the cells constantly undergoing division. natural size. especially in length. 40). about % The cells in this region increase in size. where the newly formed cells tion. entiated embryonic region gradually merges into the region of elongainch in extent. with or a thin is more less cells cubical. B FIG. 3B and C). A. large vacuoles. 3A). the same seedling 24 hours later. numerous root most layer of hairs. of the development of thicker because have slightly walls. dense embryonic nucleus The and a (Fig. showing elongation only near the tip. B. and. 40. relatively large cytoplasm. Growth of the root in length. wall. The fact that elongation is restricted to this region may readily be demonstrated by placing a row of equidistant dots along the entire length of a young root and a day or two later observing which ones have moved apart (Fig.

become complete. taken behind the root- hair zone. In addition to roothair formation. The root-hair zone extends back only a short distance and gradually merges into the mature root. Since the latter are surrounded by films of water. the intimate contact of root hairs to enables soil particles a maximum quantity of available moisture to#be absorbed. 41. the development of strands of elongated conducting cells in Root hairs are epidermal outgrowths. the center of the root. the root hairs They have a thin become firmly united with the 42). thin-walled. soil particles (Fig. is Enlarged view of one of the root hairs ahown in Fig. begun in the A FIG. 42. seedlings in moist air (A) and in soil A showing root size. shows two distinct regions: an outer cortex and a The cortex consists of rather central vascular cylinder (Fig. When roots are grown in soil. has now cross section grown (J5). 41). relatively undiffereiitiated cells forming a . wall and a lining of cytoplasm around a large central vacuole. the most conspicuous change seen in this region FIG. The differ- entiation of tissues. B Radish hairs.VEGETATIVE ORGANS OF SEED PLANTS root tion. natural younger part of a root. X 350. large. 43). absorbing surface of the rootlet. Their presence very effectively increases the. of the root. each one representing a single cell (Fig. Structure of Mature Root.. 61 become differentiated in structure and specialized in func- assuming their mature characteristics. 39.

X 30. The cells specialized for vascular cylinder of the mature root consists mainly of the conduction of fluids. As already noted. enlarged view The cortex is of the latter. in the young root. . parenchyma is a rather loose tissue. characterized by conspicuous intercellular spaces. Frequently. X 150. showing the relation of xylem to phloem. internally by the endodermis.62 FUNDAMENTALS OF BIOLOGY kind of tissue known as parenchyma. diagrammatic cross section of a buttercup root (Ranunculus acris). 43. B A. as here. The epidermis is tissue consisting of layers of small. The cortex is bounded externally by an epidermis. the layer from which. bounded externally by the epidermis. the root hairs arise. B. cubical cells that soon replaced by cork. showing an outer cortex arid a central vascular cylinder. FIG. dead. a form a tough protective covering.

some plants are able to make a vigorous growth above ground the following spring. beet. the roots increase in thickness through the activity of a special layer of cells called the cambium. In plants that live from year to year. Food Storage in Roots. lation of large quantities of food and water in their roots during the growing season. and many which live longer. Some plants that live through only one growing season. while those of the phloem are thin walled and living. accumulate food in their roots. usually forming a solid central strand. a feature that can be a carrot the clearly seen by splitting open This peculiroot. Storage roots are seen in such common plants as the dandelion. arity is in marked contrast to in which branches on the stem. Storage roots also contain a large proBy the accumuportion of water commonly 80 to 90 per cent. The xylem tissue in a root is arranged like the spokes of a wheel. and these become thick and fleshy as a result. way arise FIG. Cambial activity will situated between the xylem and phloem. Branch roots always originate in the outer part of the vascular cylinder and push their way outward through the cortex. food is mostly in the form of starch. Frequently the vascular cylinder is en- closed by a specialized layer of thick-walled cells constituting the endodermis. X ^. and between its rays are small isolated strands of phloem surrounded by more or less parenchyma (Fig. Both kinds of tissues are made up of elongated cells. 44). viz. 44. Their radish. but in some. be discussed in connection with the stem. xylem and phloem.. carrot. and sweet potato (Fig. Storage roots of beet (A) and of carrot (B).VEGETATIVE ORGANS OF SEED PLANTS two kinds of conducting tissues are present in the pteridophytes and spermatophytes. but the cells of the xylem are thick walled and dead. sugar is frequently present as well. turnip. 43). .

stems and leaves of a plant constitute its shoot system. . It is apparent that the stem connects the roots with the leaves both structurally and A leafy stem is called a shoot. so are those of the shoot system related to the air and light. (7. Three types of leaf arrangement. Just as stances in solution to the functions of the root system are related to the soil.64 FUNDAMENTALS OF BIOLOGY THE STEM Stems are concerned chiefly with the bearing of leaves. alternate leaves of Cotoneaster. bud is present in the axil of every leaf. opposite leaves of Salvia. It supports the leaves (as well as the flowers and fruits) and conducts sub- A B C A FIG. In each case one leaf does not stand directly over another so as to shade it. A. one-half natural size. and it is The stem to this fact that they owe their distinctive features. and from them. B. whorled leaves of Abelia. has two primary functions: support and conduction. 45. while all of the functionally.

Buds and Branches. . 45). 40. for the chief work of leaves mands that they be freely deex- posed to the light. It is generally true that. showing stem tive State Oi Cell division. In most plants only one leaf is borne at a node. comparatively few plants the three or leaves are whorlcd. 46). imapex and two rudimentary leaves. Leaves are not scattered over a stem promiscuously but are always borne according to a definite and symmetrical arrangement The place on the (Fig. only here that At the of Salvia grahami. and it is new is leaves are formed. the youngest being nearest the stem apex ( /upper portion of . one standing opposite the other. secondary stem tips may arise in their ' > >. Stem apex of the growing stem tip A. leaves tend to be arranged on an erect shoot in such a regular manner that one leaf does not stand directly over another and thus shade it. of same. FIG. . regardless of the number of leaves at a node. there being more at a node. longitudinal tip section.VEGETATIVE ORGANS OF SEED PLANTS 65 Arrangement of Leaves. while the regions between successive nodes are known as internodes. showing the a Small COne-shaped mass of development of young leaves and eral bll(is x 3 5 B enlarged view imdifferentiated cells in an ac. . While the leaves are still very small. The former constitutes an alternate arrangement. stem where a leaf is attached is called a node. The stems of most seed plants increase in length only at their tips. . but commonly there are two. the In a latter an opposite one. x 175> mediately behind which there arise minute lateral outgrowths that develop into leaves.

. showing the development of branches from lateral buds. and by its activity the stem increases in length. bud "opens" simply by the elongation of its internodes A accompanied by the unfolding and enlargement of its leaves. A terminal bud is present at the upper end of each shoot. The portion which is of the shoot just described constitutes a bud. B young (. and a lateral bud in the axil of jeach leaf. 46 clearly shows. thus merely an undeveloped or embryonic shoot a A FKJ. attached to the stem. A structure having the capacity of becoming a mature leafy stem. one-half natural size. FUNDAMENTALS OF BIOLOGY as Fig. 47.66 axils. The bud situated at the upper end of a main shoot is called the terminal bud. or these may not appear until The axil is the upper angle formed where the leaf is later.A) and an older (B) shoot of Cotoneaster.

48. woody plants of colder regions. -Leniicel however. It is apparent from the preceding account that the chief external difference between stems and roots arises from the fact that stems bear leaves FIG. and type of branching. The general aspect of a plant. Although the stems of most plants grow erect and support their own weight. remain dormant over the winter. or a resinous varnish. on the other hand. In nearly is -Lateral bud such winter buds. which develop into branch shoots. natural size.VEGETATIVE ORGANS OF SEED PLANTS The secondary buds. and expand during the following spring. branch indiscriminately and do not bear leaves. 67 arising in the leaf axils. develop into branch shoots and because of their position are called lateral or axillary buds (Fig. Twig of shag- bark hickory (Carya ovata) in winter condition. or may persist for ^ a longer or shorter period. growth of the shoot being here more or less the In most of a continuous process. In nearly all herbs and WSP\ many Terminal bud in woody plants of warm regions. resting buds form arid growth In such cases buds form periodic. dropping off when the bud opens and leaving charall leaf scar the lower leaf rudiments acteristic scars on the stem (Fig. Roots. during the growing season. 47). duration. The leaf in whose axil a bud is formed may fall off the stem before the branch it shoot develops. Habit. develop as scales that closely overlap and protect the delicate parts within. wax. appear in the leaf axils. those of . and that buds. is deter- mined largely by such features of the shoot system as its growth direction. according to a regular arrangement. 48). -Scale scars The protective function of bud scales is often augmented by the presence of hairs. the cells at the apex of the stem tip may continue to divide and to produce new leaves during the entire growing season. called its habit. as they are called.

. Fio. bulb of onion cut through tho \^\B. resembling storage roots in this respect. Some plants have underground stems. either twining about a support. 49. Types X of underground stems. rhizome of Solomon's seal. and bulbs 1 (Fig. tendrils. The food is stored principally in the scales or in the leaf bases. 49). Others have developed the climbing habit. glory. center. such as rhizomes.68 vines are too FUNDAMENTALS OF weak to do so. or climbing by means of anchoring roots. A. X **j. 1 A bulb is really an enlarged fleshy underground bud consisting of a very short stem covered with either thickened scales or thickened leaf bases. tubers. X %\ C\ tuber of potato. such as cucumbers. ttlOLOGY vines. These serve as food-storage organs. like a morning- Some creep or trail along the ground. or hooks.

it is year. uribraiiched. the branches growing out Such trees have a conical form. Its subterranean parts accumulate food during the first growing season and live over the winter. plants are said to be either herbaceous or woody. the main stem continues to the top as a straight vertical shaft. The manner in which a plant branches is an important factor In some plants the main stem is bud being the only one that expands.VEGETATIVE ORGANS OF SEED PLANTS 69 According to the duration of their aerial parts. The first is chargymnosperms and of those angiosperms whose seeds have two cotyledons (dicotyledons). Based on their internal structure. the a plant whose entire shoot The beet and is carrot are familiar biennials (Fig. A perennial a plant that lives for more than two years. but is seen in the palms. of seed plants belong to acteristic of all . goldenrod. canna. Some herbs and all woody plants are perennials. This type of branching. Stem Structure. spruce. produces seed. designated as excurrent. but the upper portion of the plant commonly dies back to the ground in the autumn. the main stem soon gives rise to large branches and does not continue to the top. An herb is above-ground parts (ordinarily comprising system) die at the end of the growing season. Although rather common among herbs. 44). the terminal is said to be columnar. A herbaceous perennial is a plant whose underground parts live from year to year. Most herbs are annuals. rhubarb. type of branching. arises from the vigorous development of lateral buds. In such trees as elm and oak. In many herbs and in such trees as the pine. tulip. the stems two general types. whose habit in determining its habit. the terminal bud generally not expanding at all. results from the persistent development of the terminal buds. and Carolina poplar. This horizontally from it. and many others. and completes its life cycle. Such trees have a spreading crown. Wellknown plants of this sort are asparagus. an unbranched shoot system is rare among trees. dahlia. fir. The next year the plant sends up a new shoot. while woody plants shrubs and trees have aerial stems that live from year to If the entire plant lives through but one summer. The second and less common but more advanced type is found among those angiosperms whose seeds have one cotyledon (monocotyledons). A biennial is an called an annual. gladiolus. iris. herb that lives through two growing seasons. but whose aerial portion dies at the end of each growing season. called deliquescent.

2. . 50. X 10. 1. 3. and the formation of cork.70 FUNDAMENTALS OF BIOLOGY Cortex Mechanical* tissue Phloem Cambium Mechanical* tissue Annual rings B Vascular rays Fio. successive layers of wood (xylem) formed during the three growing seasons of the stem's existence. showing increase in the amount of xylem and phloem. Diagrammatic cross section of a one-year-old (A) and of a three-yearold stem (B) of box elder (Acer negundo).

the same kind of tissue that occurs The pith is colorless. Water with its dissolved soil salts passes upward through the xylem tissue from the roots to the leaves. 51). and none at all in the autumn and winter. Both the cortex and pith are composed chiefly of parenchyma. is generally a continuous tube. Traversing number of narrow plates of paren- chyma cells called vascular rays. constituting an annual ring. cular rays often store food and aid in the radial movement of substances in the stem. but nearly always lack chloroplasts. the cortical cells contain chloroplasts. 504). but the former also gives strength to the stem. the number of larger which indicates the age of the stem. As in a young root. Epidermal cells are living. During the growing season. giving rise to new vascular tissues to new xylem outside the old xylem. 5(L4). these extend from the pith to the The vascortex. but here the outer walls of the epidermal cells have a waxy covering. while food The vascular cylinder chiefly of in solution passes the vascular cylinder are a downward through the phloem. both xylem and phloem are conductive in function. 52A). So the line of contact between . Generally each layer is sharply delimited from the others. but surrounding the xylem there is always a narrow zone of phloem. such as that of a box elder. of a Lying between the xylem and phloem is a layer of thin-walled cells called the cambium. the cells forming the cambium are constantly undergoing division. The cambium produces xylem cells in the spring than in the summer. but the xylem persists from year to year. As in the root. This renders the epidermis waterproof. shows three general regions: (1) an outer cortex. and to new phloem inside the old phloem (Fig. Increase in Diameter. The xylem consists of a series young woody stem of concentric layers formed during successive growing seasons (Fig. checking evaporation from the underlying tissues (Fig. which typically appears as a continuous ring when seen in cross section (Fig.VEGETATIVE ORGANS OF SEEp PLANTS 71 Young Woody Stem. forming a cuticle. A cross section of a typical young woody stem. (2) a hollow vascular cylinder. but most of in the cortex of the root. the cortex is bounded externally by a layer of living cells called the epidermis. 50 B). In woody plants the old phloem gradually disappears. (3) a central pith (Fig. consisting xylem tissue. but each runs only a short distance vertically.

In Cortical Stone ceJJs parenchyma Fibrous mechanical tissue Phloem Cambium Vascular ray ern Pith FIG. its first year of those tropical regions where uniform climatic conditions prevail throughout the year. X 300. Portion of the vascular cylinder of a box elder stem in growth. showing structural details. . 51. most plants do not form annual rings.72 the FUNDAMENTALS OF BIOLOGY summer wood of one season and the spring wood of the next marks the boundary between successive annual rings.

of The tissues designated as xylem and in pterido- phloem are not uniform. older stem. the latter the A. just beneath the epidermis. cc. secondary c. but are elements. wood. cross section of hortorum). Development of cork in tho stem of garden geranium (Pdargonium - ^ . through which communication is maintained between the atmosphere green cells and of the the living cortex. epidermis. cork tissue consists of cells layers of small that lack protoplasmic contents when mature and have cell walls cubical slightly thickened with a fatlike substance that renders them waterproof (Fig. differentiated into several kinds In practically all gymnosperms. cork cambium. The sapheart- wood. The cork contains numerous openings called lenticds. Conduction occurs only in the outer (younger) portion of the wood. and because the pith always remains relasmall. Conducting Tissues. t cortex. .VEGETATIVE ORGANS OF SEED PLANTS As the vascular tissues increase in 73 amount through cambial activity. After a number of years the cortex disappears. . an old woody really consists of only the following regions: outer bark tively stem rf> c (cork). pc primary cortex. <?. outer portion of young stem. former constitutes the tion FlG 52. c. cork. 48). inner bark (phloem and cambium). forming a protective It is produced by a cambium. X 150. Lenticels can readily be seen by examining the bark of almost any woody twig (Fig. B. and wood (xylem). the central (older) porcarrying no sap. as . cork develops outside the cortex. which appears external covering. 52). As in the root.

The most important of these are vessels long tubes of large diameter A FIG. and pitted. The most important elements in the phloem are sieve tubes. X 250. These are distinguishable on the basis of the character of the thickenings on their walls (Fig. so called because their end walls. annular or ringed. elongated. elongated cells. the xylem is made up of tracheids slender. while other xylem elements occur in mature stems.74 FUNDAMENTALS OF BIOLOGY phytes. and often side walls as well. 53). are perforated like a salt shaker. tracheids sperms. annular (B). by a breaking down of their Three chief types of vessels are found: spiral. 53. B Spiral (4). that arise from a fusion of tracheids cross walls. In angioliving contents. . thick-walled cells generally pointed at either end and without Their cell walls are usually pitted. and pitted vessels (C) from stem of castor bean (Ricinus communis) as seen in longitudinal section. Like tracheids. they are also but retain their protoplasm and thin walls. are frequently present only in young stems.

Mechanical Tissues.


of mechanical tissues


The function

They are give rigidity, toughness, or hardness to plant organs. especially prominent where there is a weak development of
xylem, as in many herbaceous stems. They have nothing to do with conduction. Fibers are long dead cells with pointed ends and uniformly thickened walls; they may occur in the cortex or
in the vascular cylinder.



have very thick walls but are

macrophylla)', a

Cross section of a young stem of Dutchman's pipe (Aristolochia woody vine, showing a vascular cylinder composed of isolated vascular bundles separated by wide extensions of the pith, X 20.


Figure 51 shows both short; they often occur in isolated groups. and stone cells in the region between the phloem and cortex.

Among dicotyledons the conducting reduced in amount as compared with those of woody stems. This reduction may be a result of either diminished activity of the cambium or the breaking up of the
tissues of herbs are greatly

Herbs and Woody Vines.

between which are wide extensions

vascular cylinder into separate strands called vascular bundles, of the pith. The same reduc-

tion of conducting tissues occurs in woody stems that climb In stems of this type, the cambium may connect (Fig. 54).

the vascular bundles or not, but where




often produces




only parenchyma between them. In many cases, however, the later produces vascular tissues between the bundles, so that a continuous vascular cylinder is eventually formed.

Stems of Monocotyledons. In angiosperms whose seeds have only one cotyledon, such as grasses, lilies, palms, and orchids, the stem has separate vascular bundles like the stem shown

but instead of being arranged in the form of a hollow cylinder they are scattered irregularly through the stem (Fig. 55).
in Fig. 54,

FIG. 55.

Cross section of a young corn stem (Zea mays), showing scattered
vascular bundles,



A cornstalk, an asparagus stem, or a piece of sugar cane would serve as an excellent example of this type of stem. Between the scattered vascular bundles is parenchyma. Since no vascular

formed, there can be no distinction between pith and Each vascular bundle consists of a small group of xylem and phloem cells. The stems of nearly all monocotyledons lack a cambium, and consequently the vascular tissues, once formed,

do not increase in amount.

by growth

of the

Increase in diameter takes place parenchyma between the vascular




Leaves are typically thin expanded organs that arise as outgrowths from a stem tip. They are the most conspicuous members of the vegetative plant .body and carry on the major part of the work of nutrition. Their primary function is This photosynthesis, the manufacture of carbohydrate food.

FIG. 56.



FIG. 67.




Leaf blade of climbing (Ficus pumila) which has been

its stem, showing blade (/>), petiole

and stipules (), one-half natu-

"skeletonized" by removal of the green tissue, natural size. The smallest veinlets may be seen with the aid of a magnifying lens.

ral size.

requires that they be structurally adapted to the display of green tissue to the light.

External Features.


flat portion,



Most loaves are differentiated into a the blade, and a slender leafstalk or petiole blade is the more essential part of the leaf,

the petiole merely supporting the blade and placing it in a favorable position with reference to the light. In fact, the leaves


plants do not have a petiole, the blade then being



attached directly to the stem and constituting the entire leal. Some leaves possess a pair of stipules small, scale-like appendages formed at the base of the leaf, one on each side.

noteworthy feature of leaves is the elaborate system of veins, which extends from the petiole to all parts of the blade
Veins represent a continuation of the vascular system (Fig. 57). of the root and stem and so are channels for the transport of


water and dissolved substances that pass into and out of the



larger veins of most leaves are conspicuous, especially on the lower

also give the leaf

some degree

of rigidity.


FIG. 58.

Divided leaves of horse chestnut (A) and of rose


one-half natural

side of the blade, but the smaller ones cannot be seen without the In net-veined leaves, the larger aid of a strong magnifying lens.
freely in the green tissue (Fig. 57).

veins give rise to smaller and smaller branches, ultimately ending In parallel-veined leaves,

on the other hand, the principal veins (and often the only ones) run parallel to one another, generally from the base of the blade
to the apex, as in a blade of grass.

Net-veined leaves occur


dicotyledons, parallel-veined leaves mainly


monocotyledons. In size, leaves vary from minute scales to those as large as in the bananas and palms. In form, some leaves are more or less
circular, as in the geranium and nasturtium; others are long and narrow, like a grass blade or a pine needle; but most leaves are



of some intermediate shape. The leaf margin may be smooth and even, or variously toothed, notched, or lobed. In Some cases

the lobes are so deep that the blade


divided into separate

Leaves also show great variation in thickness, leaflets (Fig. 58). character of the surface, arrangement of the principal texture, and other features. veins,
"fbtfisade parenchymotX *

Upper epidermis

, \Mesophytt ^Spongy parenchyma]


FIG. 59.

Intercellular spaces Stomafa^ Cross section of a leaf of Japanese privet (Ligii strum japonicum) X 250. In the vascular bundle (vein) the xylem lies above the phloem.

Lower epidermis

Internal Structure. cross section of a typical leaf shows three kinds of tissues: (1) an outer colorless epidermis; (2) green tissue, called mesophyll, comprising the bulk of the internal portion; (3) veins, or vascular bundles, passing through the mesophyll Typically a single layer, of living epidermal cells, (Fig. 59).


lacking chloroplasts, As in a of the leaf.


young stem,

seen on both the upper and lower surface their walls are covered with a




This makes the epideposit that forms a cuticle. impermeable to water and gases, preventing exces-


from the inner

sive loss of water

Here and there are

peculiar Openings called stomata, which in horizontal leaves are They usually entirely or largely confined to the lower surface.
are very numerous, there being square inch.


100,000 or more to the

Stomata may easily be studied by stripping off a piece of lower epidermis from a leaf of a geranium or a lily and examining it under the microscope (Fig. 60). A stoma consists of a slit-like opening bounded by a pair of guard cells. These differ from the
other epidermal


having chloroplasts.




FIG. 60. Portion of lower epidermis removed from a geranium leaf (Pelargonium), showing five stomata scattered among the ordinary epidermal cells, the latter having wavy outlines, X 500. Each stoma consists of a slit-like opening

bounded by a pair





the stomata that gases pass from the atmosphere into the leaf In many leaves the size of the stomatal opening vice versa.

can be altered by a change in the shape of the guard cells, thus Stomata are also present in affecting the rate of gas exchange. the epidermis of young stems. The mesophyll is the great food-manufacturing region of the It is typically differentiated into two kinds of tissues: leaf. palisade parenchyma and spongy parenchyma, the former usually

forming a single layer beneath the upper epidermis. The two kinds of parenchyma differ from each other in the form and arrangement of the cells, but both have chloroplasts. Palisade

are rather compactly arranged

angles to the surface of the leaf.

The spongy

and are elongated at right tissue is composed

of cells loosely arranged, rather irregular in form,
air passages


and with large them between known as interintercommunicating cellular These communicate with the atmosphere spaces. through the stomata, and provide for a movement of gases

throughout the


Each vein

consists of a single vascular bundle in

which the

xylem occurs on the upper

the phloem underneath. Around each bundle there is usually a bundle sheath composed of colorless parenchyma cells, its thickness being proportionate to the size
of the vein.

Often mechanical tissue

of large veins, its

present in the vicinity function being to give rigidity to the leaf.





The reproductive features of spermatophytes, as compared with those of the lower plants, are very striking. Complications have arisen by the development of a number of modifications incident to the formation, protection, and dissemination of seeds, and for this reason the structures associated with reproduction Seeds are are very different from those of the lower plants. both asexual and their formation involving complex organs, In all spermatophytes the vegetative body, sexual reproduction. with its roots, stems, and leaves, is a sporophyte, arising from a zygote and in turn producing spores. Moreover, all seed
a feature which
plants are heterosporous, bearing microspores and megaspores, In fact, it is is rare among pteridophytes. heterospory which makes seed formation possible. Seeds are

not organs of reproduction, but of dissemination, while fruits merely protect the seeds and frequently assist in their dispersal. The reproductive organs of the sporophyte are found in the cones The latter of gymnosperms, and in the flower of angiosperms. will be discussed first.


A flower is a specialized shoot whose appearance is necessary to the ultimate production of seeds. All flowers arise from buds,
as in the case of vegetative shoots and, similarly, may be either terminal or lateral in position that is, a flower may be formed at

the end of a stem or in the axil of a leaf or scale. A cluster of flowers is called an inflorescence, and of these there are many

A longitudinal section of a very young flower bud reveals kinds. the fact that the floral parts originate as lateral outgrowths from a central axis in the same way that leaves arise from a vegetative
stem tip. In such primitive flowers as those of the magnolia and buttercup, the central axis is somewhat elongated, as in a
vegetative bud.

In the great majority of flowers, however, the



axis remains short, forming what is known as the receptacle, and upon this the floral parts are borne in a succession of whorls.

Flower and fruit of orange (Citrus sinensis). A, flowering shoot, B, single flower with two of the petals and several stamens cut away, X 2; C, cross section of the ovary, X 3 Z>, early stage in development of fruit, X 1 p, petal; s, sepal; at, stamen; rec, receptacle; ow, ovary wall; o, ovules; sti, stigma; The ovary, style, and stigma constitute the pistil. sty, style; ov, ovary.
FIG. 61.




Floral Parts.


typical flower consists of four sets of parts

symmetrically arranged with reference to one another (Fig. 61). The outermost set, constituting the calyx, is composed of indi-

The third floral set comprises the stamens. in some plants two kinds of flowers are produced. In other flowers the ovary is sunken in the receptacle. A stamen is generally a club-shaped organ conThe sisting of a stalk or filament supporting a terminal anther. known In other flowers there may entirely separate from one another. Although most flowers contain both stamens and pistil. sympetalous and a snapdragon (Fig. constitute the perianth. a petunia (Fig. made up of Most commonly the petals are large and conindividual petals. be several simple pistils A simple pistil is often is composed of individual In some flowers the ovary is superior. may cither be separate and dis- tinct from one another (choripetalous) or more or less united to form a tube (sympetalous) Moreover. being made up of two or more simple pistils more or less united. This with hair or with a sticky secretion. QIB). which in the two chambers. These contain which are liberated when the anther is numerous is pollen grains. taken collectively. and irregular. being either white or of some color other than green. green. and a slender stalk-like style A as ovules. . usually its or otherwise modified and often is covered enlarged tip. latter contains sporangia. but in most cases it is compound. ripe (Fig. inside of and entirely free from the other floral parts (Fig. The occurring just inside the calyx. 63).84 FUNDAMENTALS OF BIOLOGY vidual sepals. a nasturtium (Fig. all the petals may be alike in size and form (regular) or unlike (irregular) l The calyx and corolla. spicuous. 61 B). Typically this is portion of the style. The two kinds may be borne on the same inferior. is the corolla. In some flowers the pistil is simple. being situated at the top of the receptacle. 202B). which occur inside the corolla. or 1 A trillium (Fig. called a carpel. 38B) is choripetalous and regular. Ordinarily these are small. choripetalous regular. or more small bodies is one The ovary a hollow organ enclosing arising is it composed from of a lower bulbous ovary (Fig. ^ach representing a pair of fused young anther are distinct. so that the other floral parts appear to arise from its summit. . 200). leaf-like parts that enclose and protect the other floral parts in the bud. 61 C). each of which is an incipient seed (Fig. sympetalous and irregular. and so a compound pistil carpels. is termed the stigma. one lacking stamens and the other a pistil. next set. 198). The petals. as well as the sepals. In the center of the flower the pistil. .

. is accomplished in various ways. and castor bean. others neither corolla Pollination. depending on the be transferred from a stamen to the same flower (self-pollination) or to the pistil of . or on separate plants. This act of called.REPRODUCTION IN SEED PLANTS 85 plant. In pollen must first be transferred from a stamen to the pistil. as it is deposited on the stigma. as in corn. corolla. angiosperms pollen pollination. pistil Pollen may insects. Embryo sac Integuments Stalk Fio. by kind of flower. etc. nor calyx. Some flowers have no as in the willow. poplar. 62. In order that the ovules may develop into seeds. X 250. of the by wind. of ovule Micropyfe Longitudinal section of a lily ovule (Liliurn) containing a mature embryo sac. squash. such is as by direct contact. and hemp.

X 30. The Ovule and Embryo Sac.86 FUNDAMENTALS OF BIOLOGY (cross- another flower of either the same plant or another plant pollination). 62). The outer portion consists of one or two integuments which do not completely invest the ovule. 63. (From " Chamberlain. has a characteristic structure essentially the same in all Cross section of a mature anther of lily (Lilium) just as the pollen FIG. discussed in greater length in a later The stamens and pistil are called the essential organs of the flower. each contain"(Fig. in seed formation. 62). grains (microspores that have begun to germinate) are being shed. and one enlarges to form the embryo sac. seeds may be organs. The ovule. because they are primarily involved sepals and petals are known as accessory In fact. It should be noted that the functional megaspore does not leave the ovule to germinate. a linear row of four megaspores arises.") seed plants (Fig. but leave a narrow passageway at one end called the micropyle. . cells of the young ovule. produced whether a perianth is present or not. Elements of Plant Science. 296-299). Ordinarily three of these degenerate. Within the . for it is this feature that makes possible the subsequent formation of a seed. It is attached to the inside of the ovary by means of a short stalk. since they play an incidental part. which is really a sporangium. Pollination is chapter (see pp. ing a nucleus the middle one of the three at the micropylar end is . The embryo sac of angiosperms is organized in a peculiar way At each end are three small naked cells.

While the tube is rise to developing. the single nucleus divides to form a tube nucleus and a Generative nucleus Tube nucleus Inner wall Outer wall FIG. only one pollen tube enters each of the ovules present in the ovary. before the pollen grains are liberated from the anther. 64. puts long After reaching the cavity of the ovary. arisen by three successive divisions from the nucleus of the megaspore (Fig. The generative nucleus is surrounded by a small amount of cytoplasm 750.REPRODUCTION IN SEED PLANTS the egg. As a rule. but sometimes earlier. are formed in groups of four. which may or may not be organized as cells. 654 and B). 87 In the center of the embryo sac are two nuclei. like the megaspores. 64). Section of a pollen grain of a lily (Lilium) in the shedding condition. The pollen tube penetrates the embryo it sac and discharges into the -two male cells. At first each consists of a single uninucleate cell with a rather heavy. called These eight nuclei have polar nuclei. this pollen tube grows cell (Fig. in contact with each other. two-layered cell wall. Although many pollen grains may germinate on the stigma. X generative nucleus. as the case . The tube nucleus finally disintegrates. This it then enters through the micropyle. but all of them mature. or male nuclei. the latter commonly being organized as a small Upon reaching the stigma. each pollen grain forth a tube that grows down the inside of the style. naked Fertilization. The pollen grains are formed in sporangia within the anthers (Fig. and so is organized as a naked cell. Pollen and Pollen Tube. the generative nucleus gives two male nuclei. 63). and a plasma membrane. naked along the ovary wall until it reaches one of the ovules. being in reality a microspore. The microspores.

corolla. dropped off. the ovary wall has formed the wall of the fruit. A. 65C) . 65. and often the sepals do likewise. One male nucleus is shown uniting with the egg. ripe fruit. joining FIG. the zygote has given rise to the embryo. wither and drop off. the endosperm nucleus to the abundant endosperm tissue (shown in (From Sinnott. The calyx and corolla are shown in solid black. they are non-ciliated. The may The male cells other undergoes a unique performance in angiosperms. The other cells in the embryo sac are functionless Following fertilization. has taken place. Diagrams illustrating the formation of the fruit and seed. bud. are really sperms. and the pollen tube from one of them has penetrated the ovule and discharged its two male nuclei into the embryo sac. and stamens have the two polar nuclei. One of the male nuclei penetrates the egg. "Botany. and embryo dotted. B. C. the outer portion of the ovule has formed the testa of the seed. the petals disappear. fully opened flower. the ovule containing a single megaspore. young the ovary wall and style lined. bud ready to open. and soon and stamens The ovary . The ovary contains a single ovule. Two pollen grains have germinated. but in all seed plants except a few primitive gymnosperms. the other with The calyx. testa.88 FUNDAMENTALS OF BIOLOGY be. The megaspore has produced an embryo sac with its egg cell below and the two polar nuclei The anthers have burst and pollination in the center. then all three nuclei fuse to form the primary endosperm nucleus (Fig. the ovule. D.") the two polar nuclei in the center of the embryo sac. Principles and white) which surrounds the embryo. forming a zygote. Problems.

The Fruit. thus completing the life cycle. but in many other angiosperms the endosperm persists. gives rise to the sporophyte. or rudimentary plant. The relation of the reproductive structures of spermatophytes to those of the lower plant groups is somewhat difficult to understand unless a more extensive study is made than has been outlined in the present should be noted. and the ovule becomes a seed (Fig. two kinds of gametophytes occur. producing two kinds of spores: microspores (small) and megaspores (large). Embryo and Endosperm. resulting from the fusion of a male with a female cell. The fertilized it egg. After fertilization has taken place and the ovules are ripening into seeds. colorless. embryo does not proceed very far in its development before it goes into a dormant state. however. arising from a microspore. 89 and the contained ovules are transformed Alternation of Generations in Seed Plants. developed by a megaspore. but in some fruit consists essentially of a ripened ovary. and from there arises. and entirely dependent for nourishment upon the sporophyte. zygote. the ovary itself enlarges and undergoes other changes that result in the formation of a fruit (Figs. while the embryo sac. It of seed plants only one is functional. although four megaspores arise within each ovule. The zygote. 61 and 65D). The microspores are shed. chapter. an embryo. or germinates within the ovule. by the processes of growth. into seeds. The nucleus. so that the mat is entirely absorbed by the developing lire seed lias none. At the same time the primary endosperm originating by a triple nuclear fusion. The microspores arise in large numbers within the anther. but the megaspores are not. cases associated parts also enter into its formation. both being formed in sporangia. A . a nutritive tissue that surrounds the embryo and contains stored food. that the vegetative body is a sporophyte. 65Z)). Meanwhile the integuments of the ovule harden to form a tough outer covering. The contents of the pollen tube. obscure. gamctophyte. In some angiosperms the food stored in the endosperm embryo. and. represent a greatly reduced male gamctophyte.REPRODUCTION IN SEED PLANTS enlarges to form a fruit. gives rise to endosperm. is really a female Both kinds of gamctophytes are small. As in the heterosporous pteridophytes.

which of small ripened ovaries are embedded in the surface of a fleshy At maturity fruits may be fleshy. as in the olive. lower view of same. the ripened ovary wall forms two distinct portions: an outer fleshy and an inner stony one. C. like such as the cherry. which grows up around the ovary. 66. developed under man's influence. E. a number In the strawberry. Pine cones and cone scales. D. X 4. a carpellate cone. becoming more or less fruits.90 If FUNDAMENTALS OF BIOLOGY only the ovary is involved. peach. X !% bean or pea pod. Fruits one seeded. but if other For example. the latter enclosing a single seed. Sometimes a group of fruits ripen together. as in the blackberry " Seedless" fruits are unnatural products raspberry. fruits may split open. features of The reproductive gymnosperms and angiosperms are similar in many . B. an accessory fruit is formed. or many seeded. or thick skinned. or plum. parts are included. dry as in the peanut. In stone and consolidated to form an aggregate fruit. like an acorn." the fleshy part of the fruit developing chiefly from the receptacle. a true fruit results. as a A ^ D Mf E FIG. like a melon. Reproductive Features of Gymnosperms. like a peach. When receptacle. or dry. orange. side view of a single stamen. longitudinal section of a staminate cone. each stamen hearing a pair of microsporangia on its lower side. as in the tomato or grape. A. in the apple the ovary forms the inner portion of the "core. may be Some an fruits are thin skinned. also is an accessory fruit. a single carpel bearing a pair of megasporangia (ovules) on its upper face. or remain closed.

flowers are thought to have been derived from cones. Moreover. 66). although the cones that they bear correspond to the flowers of angiosperms. 68). one-half natural size. 67. In fact. bear two kinds of cones one made up of stamens. pod of Lima bean split open to show the enclosed seeds. 91 but different in others. it produces several archegonia at the micropylar end and thus is a Each archegonium contains a single true female gametophyte. 0fc B Gyrnnosperm and angiosperm compared. B. Both the stamens and carpels : are scale-like and rather different from the corresponding parts A FIG. and the other of carpels. Gym- nosperms. In gymnosperms the megaspore gives rise to a mass of tissue within the ovule that corresponds to the embryo sac of angio- sperms but obviously is more primitive (Fig. These cone scales are borne on an a condition retained by certain primitive flowers. 67). such as a pine. in angiosperms (Fig. cone scale of sugar pine bearing two naked seeds at base of upper surface. and hence the seeds as well. the pollen egg. as already noted. No perianth is present. Since the ovules are not enclosed in an ovary. grains came in direct contact with them and so have only a . A. elongated axis. flowers. The ovules. noted.REPRODUCTION IN SEED PLANTS respects. are not enclosed in an ovary but occur freely exposed on the flat face of a carpel (Fig. gymnosperms being in general like will more primitive and consequently more Some of these important differences Gymnosperms do not have true now be pteridophytes. natural size.

embryo tion occurs as in angiosperms. in the former group appearing before fertilization. which urn the most primitive existing gymnosperms. As a rule only one egg in each ovule gives rise eventually to a fully developed embryo. in the latter after fertilization. but in addition to the tube nucleus there are several functionless antheridium is present. The seed of gymnosperms is essentially similar to that of angiosperms.92 FUNDAMENTALS OF BIOLOGY The short distance to travel in order to reach the archegonia. No Femafe 4 gcrmefophyfe brchegonium Po/len tube Infegumenf Micropyle Longitudinal section of a pine ovule (Pinuts) containing a mature FlO. produces as a rule only two male cells. X 15. which is always present. is the presence of swimming sperms. while " differarises in two in both called endosperm" cases. except that the endosperm. tissue in the seeds of gymnosperms and of angiosperms. distinctly ent ways. 68. but there is no triple fusioii of nuclei to form endosperm. tube and in the male gamctophytc. contained developed pollen from a pollen grain (microspore). Fertilizacells. sac. . A remarkable feature of the eycads. Thus the food-storage represents female gametophyte tissue.

its growth. of the hypocotyl just : Dormancy. The dormant condition of the embryo enables the seed to become detached from the plant that produced it and to remain viable until conditions are favorable for its renewed Thus seeds are organs of dissemination. a stem that gives rise to a root at its lower end. 69). All seeds have an embryo and nearly all have a testa. to or wa. morning-glory. Seeds are really not organs of reproduction. and the monocotyledons with one. (3) one or more leaf -like cotyledons. Because they are organs of is As already dissemination. which is the new plant.veined leaves and stems in which the woody tissue forms a hollow cylinder. buckwheat. etc. but the embryo. plant is bean. An interesting correlation is seen in the fact that dicotyledons have net. sunflower. have two all On this basis but angiosperms the dicotyledons. enabling plants become distributed over a wide territory. the seed is A resulted in the endosperm. When a seed germinates. and generally also a special food-storage region. and squash the endosperm is entirely absorbed by the embryo during its early development. by animals. (2) short very a minute bud arising from the upper end of the the plumule. angiosperms are divided into two great groups with two cotyledons. a new not produced. seeds pass into a state of dorduring the ripening process. The seeds many gymnosperms either or one. while in corn. 295-296). (see pp. They are commonly adapted to dispersal by some special means. wheat. In the pea. attached to the below the plumule (Fig.REPRODUCTION IN SEED PLANTS THE SEED 93 a ripened ovule in which antecedent processes have development of a dormant embryo. consists of three main parts: (1) the hypocotyl. upper end of have a number of cotyledons.ter. as by currents of air growth. while monocotyledons have mostly parallelveined leaves and stems with woody tissue in scattered bundles. the arresting of vital activities during dispersal . but when ripe many lack endosperm. renews simply Seed Structure. and castor bean the endosperm remains in the mature seed and is not used by the embryo until germination begins. for reproduction takes place when fertilization of an egg results in the formation of a zygote from which the embryo develops. The embryo hypocotyl. an outer seed coat or testa. mancy stated.

X 1. being commonly 10 to 12 per cent by weight. X 3. a seed without large cotyledons. viable for more than 3 or 4 years after ripening. are extremely rare. numerous experiments have proved that such seeds will not grow. growth will proceed. if the embryo is still alive. If for several months before germination can external conditions are then favorable. and so the grain is really a orie- eeeded fruit. Accompanying this water loss. Such cases. amount is small. notably of certain legumes. retain their viability for a long time. yet the seeds of some plants. but if not. however. Although all mature seeds contain some water. A. 69. B. the food being stored in the two of the embryo. . germination begins by the resumption of growth and other activities within the seed. Conditions for Germination. will germinate even if more than a hundred years old. the seeds of most plants will not retain their Few seeds will remain viability for more than a year or two. must remain dormant occur. Most ripe seeds will not sprout immediately upon ripening. important chemical changes occur. the resulting in The power if of all vital processes. Stories of viable wheat being taken from ancient Egyptian tombs have no scientific foundation. a Lima bean laid open to show the three parts There is no endosperm. in fact. longitudinal section of a grain of corn. Dormancy is brought about primarily by the withdrawal of water. of After a longer or shorter period dormancy. but ffypocofyf plumule Hull Tesfa "otyledons Endosperm Cofyledon Plumule Hypocofyl Roof fip FIG. Some seeds are able to grow as soon as ripe. Seed structure. endosperm and with a monocotyledonous embryo. but if conditions are unfavorable for growth.94 FUNDAMENTALS OF BIOLOGY a necessity. of time that seeds are capable of retaining their length varies of germination greatly % The seeds of willows die an almost complete cessation they do not sprout within a few days after falling from the tree. The outer covering consists of the ovary wall fused with the testa.

r. from plumule. of the Seed- shoot developing The first external evi- dence of germination is the rupturing of the testa and the appearance of the root tip. (2) There is a decrease in the amount of is reserve food. Water and oxygen pass by osmosis through the testa into the seed. The hypocotyl does not elongate and the cotyledon remains within the seed. only if three external conditions are satisfied: (1) The temperature must be favorable. (3) There must be an adequate supply of oxygen. as can be shown by* placing some of them with a thermometer in a vacuum bottle. Heat is evolved by sprouting seeds. size. growth of the embryo cannot proceed.REPRODUCTION IN SEED PLANTS 95 This happens. showing that it being consumed by the (3) embryo. firmly anchoring the seedling. FIG. from the lower end appear. the . Development ling. acting as an absorbing organ. The primary After it is root arises formed. The root grows straight downward and sends out root hairs into the soil. Soon short lateral roots primary root. however. ones. the involving absorption of liberation of the oxygen and carbon dioxide. now becomes accelerated. half natural Successive size. 70. being manifested in three ways: (1) There is always change seeds a vigorous gas ex- between and sprouting the atmosphere. of the hypocotyl. stages in the development of a corn seedling. former The which process of respiration. If any one of these three conditions is not fulfilled. (2) Abundant moisture must be available. previously was very greatly feeble. The absorption of water softens the seed coat and causes the seed often to twice its to swell.

pea. hypocotyl. A. cotyledon. After the root has . 71). A more complex type of seedling development is seen in the common garden or kidney bean (Fig. c. By the elongation of the hypocotyl the cotyledons are pulled out of the soil. and the cotyledon or cotyledons remain in the soil. elongate. Corn. r. 70).96 hypocotyl FUNDAMENTALS OF BIOLOGY may remain short or it may undergo considerable elongation. and scarlet runner bean illustrate the first type of Here the hypocotyl does not seedling development (Fig. primary root. contributes to the growth of the seedling until the roots have established a connection with the soil and the leaves have expanded to the light. s. Successive stages in the development of a bean seedling. depending upon the type of germination. the development of the plumule being retarded until this is accomplished. 71. FIG. shoot developing from plumule. putting forth green leaves at the surface of the ground. The plumule gives rise to a shoot that pushes its way upward. natural size. The reserve food in the endosperm or cotyledons. as the case may be.

morning-glory. a root. showing vegetative propagation by means of runners. The dying away of the runners isolates the new plants that have arisen at the nodes. It marble.. testa Here the single cotyledon is pulled from the and functions as a photosynthetic organ until the first true leaves appear. the others remaining dormant. and castor bean. they wither and drop off. At first the growth uneven. VEGETATIVE PROPAGATION In many angiosperms stem. or leaf. The plumule.REPRODUCTION IN SEED PLANTS formed at the lower end to of 97 of the hypocotyl. one-half natural size. not so common among monocotyledons as the first type but is seen in the onion. resulting in the formation of a The bent hypocotyl then straightens. Note that only every other bud develops. the cotyledons out of the testa and carrying them upward into the air. the latter continues is developed. but as soon as their food supply has been exhausted. At the same time the cotyledons spread apart and turn green. now gives rise to a leafy shoot. situated between the cotyledons and up to this point relatively inactive. Strawberry. Other dicotyledons illustrating this second type of germination is are the sunflower. reproduction may be brought about by This is called vegetative propagation . grow as the root system the hypocotyl is FIG. pulling hypocotyl arch. 72.

The planting of " cuttings multiplication extensively used many cultivated plants (Fig.98 because it FUNDAMENTALS OF BIOLOGY involves merely the isolation of some vegetative part For example. onion are all means by which vegetative multiplication "slips" or is accomplished (Fig. and FIG. is a method of vegetative by gardeners in propagating Most commonly stem cuttings " are planted. illustrating the formation of roots at the lower end of a branch previously removed from a mature plant and placed in soil. stem branches called runners. some The "eyes" of a potato are rise to cases root or even leaf cuttings are nodes at which are two or more minute buds that give new plants when pieces of . but in utilized. in the iris The formation and away and isolating the new plants (Fig. which creep along the ground. 73). 49). of tubers in the common potato. Cutting of chrysanthemum. 73. horizontal of the plant body. These send out roots and buds at the nodes. the older parts later dying 72). of rhizomes and of bulbs in the tulip lily-of-the-valley. are produced. in the strawberry.

flowers. is a . especially of fruit trees. 49C). such as the navel orange. and seeds that it plant. of would have produced had it not been removed from its own The practices of budding and grafting provide a means propagating desirable races of plants. fruits. In the practice and grafting. same kinds of leaves. it is healed. the use of some method of vegetative propagation obviously necessitv. The detached member becomes part on which it has been inserted. a detached bud or twig of one plant is inserted into an incision made in another in such a way that the of budding two will grow with wax until of the plant The wound is then wrapped or covered together. but always produces.REPRODUCTION IN SEED PLANTS 99 the tuber are put into the ground (Fig. In the case of seedless varieties. the of course.

Inorganic substances cannot do this. fats. All processes in plants concerned with the manufacture and utilization of food. and that plants which lack chlorophyll are unable to carry on this important function.CHAPTER VII METABOLISM AND IRRITABILITY IN PLANTS The term metabolism is applied to all processes in plants and animals that are concerned either with the building up or breakIn other words. such as the Indian pipe. while those that subsist on other living organisms are parasites. Independent and Dependent Plants. but raw materials from which foods are constructed in the plant body. These are not foods themselves. sugars. dodder. Foods are organic substances capable of yielding both energy an i formative material that living things may employ in carrying on their vital processes. 226). and proteins. whether plant or animal. they etc. and the elimination of waste products are phases of metabolism. Green plants are said to be independent because they nourish themselves.). it is the totality of ing down of protoplasm. any metabolic process may be either constructive or destructive. carbon dioxide. the release of energy. These are organic substances. while dependent plants must absorb their food from some external source. depending upon whether it contributes to the building up or to the breaking down of living matter. and mistletoes Dependent plants that absorb food from (Figs. . Protoplasm. can derive energy from only three classes of substances: carbohydrates (starch. All plants are independent except the fungi and a few arigiosperms. physical and chemical changes by means of which the life of an organism is maintained. 225. It has already been pointed out that all green plants have the power of making food by photosynthesis. They build up food within their own bodies from water. dead organic matter are saprophytes. 221. and mineral salts inorganic substances that they absorb from their environment. In general. Independent plants support themselves.

Oxygen and carbon dioxide substances the leaves and enter young stems through the stomata. called humus. they stems the older enter through the lenticels. not a source of food for green plants. carbon dioxide. until finally the entire liquid is uniformly light blue. it obtains oxygen and carbon dioxide. Oxygen. minute particles of the salt escape. it adds to the mineral content of the soil and thus enriches it for green plants. and mineral salts are the only substances that a green plant absorbs from its environment. mingling of the particles of one substance with those of another . Water and its soil. After humus is completely decayed. but while it is still undccomposed it is not utilized. Then the salt is said to be dissolved. A green plant has two sources from which it derives substances essential to its existence. Organic matter in the soil. the difference between green plants on the one hand. as is evidenced by the dark-blue color assumed by the water around the crystal. Thus the food of all organisms is the same. It is only green plants that can synthesize food from inorganic materials.. and animals and dependent plants on the other. and a solution This spontaneous interof copper sulphate in water is formed. viz. Green plants are the ultimate source of all nourishment. Absorption. especially its leaves. the air and the Through its shoot system. physical process that can easily be understood. Animals and dependent plants must obtain it from an external source food that was originally formed in the body of a green plant. arising from the manner in which they obtain their food. and without them no other forms of life could exist. water. All substances enter the plant by diffusion. Through its root system it absorbs water with oxygen and small quantities of certain mineral salts The oxygen is used in respiration. indicating that a complete mixing of the two substances has taken place. is not absorbed. through the activities of saprophytes.METABOLISM AND IRRITABILITY IN PLANTS 101 alone yield nourishment. a Diffusion. That but this contains such organisms as it is nourishment is evidenced by the fact that mushrooms and earthworms can live on it. in food manufacture. If left undis- turbed. dissolved substances are taken into the roots chiefly through the root hairs. gradually the colored area becomes more and more extensive. the other dissolved in it. they alone can sustain life. When a crystal of copper sulphate is dropped into a glass of water.

that the same is is. the salt particles diffuse into the pure water..e. Although practically all organic membranes are permeable to any given one is usually permeable to some dissolved substances and impermeable to others. Diffusion exhibited only by not dissolve in water. 74. its particles. and consequently there is no movement of If the liquid is shaken. 74).102 is FUNDAMENTALS OF BIOLOGY (Fig. but the starch soon settles to the bottom. and the water a salt solution is When particles into the salt solution. from a place is of higher to one of lower concentration. called diffusion The direction of movement is always from a region where the particles of the dissolving substance are closer together to where they are farther apart. "Botany. Principles and Problems. a membrane said to be permeable if it if the substance can pass through it. With respect to any given substance. separated from pure water by a membrane through which the particles of both substances can pass. i. Osmosis. A lump of starch will FIG. (From Sinnott.") sion. Diagram illustrating the diffusion of the molecules escaping from the surface of a soluble crystal placed in a vessel of water. Osmosis is the passage of water or of dissolved substances through a membrane and thus is a special kind of diffu- soluble substances. until there are approximately number of salt particles per unit volume on one side as on the other. and impermeable water. Thus a thin piece of . This movement continues until the concentration is the same on both sides of the membrane. as if no membrane were present. a mechanical mixing occurs. cannot.

water is with- over the lower end of a thistle tube partially filled with a strong salt solution arid immersed iri a beaker of water. this A familiar illustration of is FIG. unbroken. or. An occurs is osmotic movement of water whenever a strong solution separated from a weaker one by a membrane that is permeable to the water but impermeable to the dis- solved substance. Under these conditions. in other words. into the concentrated As a result. from the less dense to the more dense solution. under certain conditions. to water and to practically all dissolved substances. but a peculiar thing happens a great deal salt passes no of tion. The direction of movement is always from the solu- tion of less to that of greater concentration of particles of dissolved substance. . practically through the membrane into the water. but any given soil salt. the solution (Fig. that water moves in the opposite direcis. soil. by osmosis that all substances enter and leave all living The cell wall that surrounds most plant cells is permeable cells. is impermeable to the sugar contained inside. if in water. the osmotic Demonstration of phenomenon seen in the swell- A movement of water. water enters. almost completely checks the free diffusion of the particles of salt. But if the swollen fruit is put into a strong syrup or a strong brine. and if the salt solution in an enclosed space. a considerable pressure develops. may enter. parchment membrane is tied ing of prunes or raisins when placed The skin of the fruit. while permeable to the water particles. 75). and thus solution and rise in its level. Water passes into the tube. placed between a salt solution and pure water. causing a dilution of the salt drawn and It is it again shrinks.METABOLISM AND IRRITABILITY IN PLANTS 103 parchment. 75. but the plasma membrane is permeable to some substances and impermeThus sugar in a root hair cannot pass into the able to others. volume of the liquid increases on one side of the membrane and decreases is on the other.

Conduction and Transpiration. This behavior is known as plasit molysis. B As a consequence. table salt. water \PIasma membrane enters because the concentration is of dissolved substances 'ellwall greater inside than outside But if placed in a the cell.walled tubes. the through the living water and its dissolved substances pass the cortex . If the cell is not left too long in the salt solution. and thus the ascending water with its dissolved substances (sap) moves through cavities in colls of the root hairs. a mass movement of water and solutes takes place upward through the xylem tissue into the stem and out into the leaves. cell is in Plasmolysis. Xylem. common water passes out of the A F'IG.104 FUNDAMENTALS OF BIOLOGY of water depends upon the total concentration of It dissolved substances on each side of the plasma membrane. being shrinks wall elastic. After entering soil by osmosis. both into the root The entrance and from If cell to cell. it will be recalled. The cell sap of the root hairs contains certain dissolved substances to which the plasma membrane is impermeable. of and of water is important to understand that the particles dissolved substances move independently of one another. Vacuole stronger solution of a substance to which the plasma membrane such as i s impermeable. the same after 5-minute immersion in a 2 per cent solution of sodium chloride. will normal appearance when put back into pure water. especially sugar. a living contact with water or a very weak solution. thick. regain its of water into the root hairs occurs. They are greatly elongated. cell. Upon reaching the vascular cylinder. 76). the plasma Demonstration of plasmolysis. away from the cell and the protoplasm forms a spherical mass within the cell cavity (Fig. B. consists almost entirely of cells that have no protoplasm. and is relatively more So an osmotic movement highly concentrated than the soil water. 76. the root if Dissolved substances also may diffuse into them the plasma membrane is permeable to and certain other conditions are favorable. A. membrane. showing the contraction of the plasma membrane caused by a withdrawal of water from the cell. X 250. normal cell of leaf of Elodea.

A geranium shoot. The giving off of water vapor from leaves is termed transpiration^ but . held in an inverted flask by means of a split cork. the plant would wilt and eventually die. Demonstration of transpiration.METABOLISM AND IRRITABILITY IN PLANTS dead cells. evaporating from the leaves. 105 osmotic Such a mass movement is much more movement through living cells would be. water that a plant absorbs from the to replace that lost into the air. the chief value of the constantly being given off into the air from leaves. condenses on the inside of the flask. If this were not constantly replaced. nearly the water that has traveled upward through the stem evaporated through the stomata. has its lower end in a bottle of water. FIG. Water vapor soil is is in fact. 77. Moisture. all is rapid than an Upon reaching the leaves.

passes into the intercellular spaces. 77). This water loss. passes from the The ends of the veinlets to the mesophyll cells.106 it is essentially FUNDAMENTALS OF BIOLOGY merely evaporation through stomata (Fig. but as yet a thoroughly It is satisfactory solution of the problem has not been reached. and as a result it continues to enter the leaf by The proportion of carbon dioxide in the atmosphere is 0. Loss of water into the air from the mesophyll cells results in a concentration of their cell sap. Photosynthesis. Carbohydrates are the products of photosynthesis and constitute the The steps in their formation will be briefly chief food of plants. The necessity of carbon dioxide to the manufacture of carbo- hydrates can be readily demonstrated by placing under a bell jar a vigorous potted plant and a small dish of caustic potash. through the conducting system. carbohydrates are made in cells containing chloroplasts. and becomes dissolved in the water that satuUtilization of this rates the cell walls of the mesophyll tissue. is unavoidable because the stomata must be kept open so that respiration and photosynthesis may go on. apparent that an enormous amount of energy is required to cause water to rise to the height of tall trees.2 per cent nitrogen. Under these condi- . a substance that absorbs carbon dioxide. only diffusion. and 20. and so water is drawn osmotiTranspiration exerts a cally from the open ends of the veins. tremendous pull on the innumerable threads of water in the xylem. and yet the particles of water in these minute threads seem to hold together.8 per cent oxygen.03 about per cent. One of the most widely accepted theories seeks to explain the ascent of sap in terms of the lifting force of transpiration and the cohesive strength of water. always a danger to the plant. as compared with 78. Physicists say that the cohesive strength of water is much greater than is commonly realized. The causes governing the upward movement of water in the plant have been extensively investigated. Carbon dioxide from the air enters the leaf through its stomata. Raw Materials. gas in photosynthesis reduces its concentration in the intercellular spaces. considered. When light falls upon a green plant. are water and carbon dioxide. The source of this energy is still an uncertain matter. hydrates are made moving upward inorganic substances from which carboSoil water.

the chloroplasts have the power of combining carbon dioxide with water in such a way that a carbohydrate is formed and free oxygen is liberated. however. Photosynthesis can proceed only in the presence of At night the process ceases. have variegated leaves. hydrates are these plants are exposed to the light. Upon testing for starch after the plant has been illuminated for several hours. 78). In the presence of light. leaf. thus demonstrating that photosynthesis is dependent upon chlorophyll. It does this.METABOLISM AND IRRITABILITY IN PLANTS tions 107 no carbohydrates are formed in the leaves. it is apparent that the presence of chlorophyll is essential to it. the great center of photoall green parts of the Certain varieties of cultivated plants. The agent in the manufacture of carbohydrates is the chloroplast a special- body containing chlorophyll. This is because the chlorolight. ized protoplasmic Although the mesophyll of the leaves is synthetic activity. A potted plant is kept in the dark for 24 Then a piece of tin foil or opaque paper is fastened to hours. necessity of light in photosynthesis can be shown by a simple experiment. As light passes through the leaf. The one of its leaves in such a light. plasts absorb radiant energy from the sunlight and employ it in the conversion of the raw materials to carbohydrates. It should be noted that apparently it is not the green pigment itself. and this is why leaves appear yellowish green. Since photosynthesis Agent. only under the influence of the chlorophyll. as in the cortex of young stems. Energy. the process goes on in plant. it is found that none has formed in the shaded from the but still has access to the portion of the Process. way that a portion of the leaf air is covered from below (Fig. is a function restricted not only to green plants but to cells containing chloroplasts. in which portions of the leaf fail to develop chlorophyll. . the red rays are principally absorbed. but to some extent the blue and violet ones also. The other rays are not absorbed but are reflected. carboformed in the green parts of the leaves but not in When the white parts. such as geraniums. but the living matter in the chloroplast that carries on photosynthesis. Other experiments have proved that green plants obtain from the air as carbon dioxide all the carbon that is used in constructing their organic compounds.

Experiment demonstrating the necessity of light in photosynthesis. the leaf was illuminated. The liberation of free oxygen during photo- 1 Actually. a piece of tin foil in which a star-shaped hole has been cut is attached to the upper side of a leaf by a light screen that admits air to the lower side but excludes light. Courtesy of Bausch and Lomb Optical Co.} and six of free oxygen (each molecule of oxygen being composed of two atoms). and finally into the carbohydrate. the equation representing only the end products of a series of intermediate reacIt seems that the water and tions that are incompletely understood. 1 The primary product of photosynthesis is gen- known as grape sugar.108 FUNDAMENTALS OF BIOLOGY process of photosynthesis : The is represented by the following equation 6CO 2 Carbon dioxide + 6H 2 - C Hi 2 O 6 6 + 6O 2 Oxygen Water Grape sugar This means that six molecules of carbon dioxide are combined with six molecules of water to produce one molecule of grape sugar . liberated as a waste product. (A. passes out of the leaf through the stomata. A B FIG. the process of photosynthesis is not so simple as this. although some may be immediately erally a simple carbohydrate used in respiration. the same leaf treated with iodine after several hours' exposure to the The dark area indicates the presence of starch. which formed only where light. J9. carbon dioxide are first decomposed by the chloroplast. . Most of the free oxygen. their constituent atotns then being recombined. 78. first into one or more simpler compounds. A.

food is FIG. 79). Ordinarily sugar is formed in a leaf more rapidly than it can be rethe plant body.METABOLISM AND IRRITABILITY IN PLANTS synthesis 109 may be observed by placing a jar of aquatic plants in sunlight (Fig. an insoluble carbohydrate. apparatus is standing in sunlight. (2) to vital oxygen in. The excess is then commonly changed to starch. thus result- The displacing the the test tube. Fats are compounds of . for it is the potential energy contained in all foods that makes them capable of sustaining life. lustrating Experiment il- conducted through the phloem. which temporarily accumulates. activities to go on. It should be recalled that. for the most part. is stored in a wound watch This storage of energy is really the most significant phase of photosynthesis. form new protoplasm (in assimilation). ing in growth. but are transformation products derived from carbohydrates. foods than carbohydrates. Utilization of Food. Food may be utilized by living cells in two ways: (1) as a source of energy (in all thus enabling respiration). after photosynthesis has stopped. Sugar constitutes the chief food of plants and is the basis of all other organic substances formed in After being made it be immediately used by the leaf cells as a source of nourishment. 79. The energy absorbed from the sunlight and utilized by the chloroplasts in bringing about the union of carbon dioxide and water is stored in the. thesis directly These are more complex They are not made by photosyn- from inorganic substances. Bubbles of oxygen are arising from the cut end of the stem of the water and are plant (ELodea) gradually water in the release of photosynthesis. or transported through the vascular system to other parts of the plant and there utilized. At night. carbohydrate just as energy spring. Formation of Fats and Proteins. the starch is reconverted to sugar and transported in solution to other parts of the plant. may moved.

it may occur either light or chlorophyll. but here they are combined in a different way. hydrogen. All the seven other elements enter the plant from the soil in the form of salts. not merely in of green parts. some protein food because protoplasm Proteins are compounds itself is largely composed of proteins. most frequently is stored in the form of . but in fats the proportion oxygen to hydrogen is much less. In carbohydrates the hydrogen and oxygen are nearly always combined in the same proportion as they occur in water. and phosphates. and phosphorus enter the plant through its root system in the form of nitrates. All organisms require four-fifths of the air consists of free nitrogen. Minute traces are potassium. calcium. None of these is used in protein synthesis but in other ways. oxygen. the same elements that are present in carbohydrates. in time and any living part of the plant. Food Storage. that is. Food made by the plant and not immediately used accumulates in various parts of the body. sulphur. Thus the ten principal elements that the plant must have in order to carry on its normal functions are carbon. as it is not place at any In fact. They can be formed in any part of the plant where carbohydrates are present. phosphorus as well. In addition to nitrogen. and oxygen. magnesium. nitrogen. while most of them some of and contain sulphur them. green plants are unable to utilize atmospheric nitrogen in the synthesis of proteins but must depend for their supply of nitrogen Although nearly The manufacture of proteins may take soil. hydrogen. and oxygen. the elements obtained from carbon dioxide and water. It should be realized that by far the greater part iron. dependent upon their which in dependent plants. sulphur. sulphur. and phosphorus. They are to which new elements are added to elaborated carbohydrates form very complex compounds. phosphorus. and of carbon. sulphates. The elements nitrogen. all green These plants require four other elements that occur in the soil. of the plant is composed of carbon. as salts. oxygen. magnesium. hydrogen. carbohydrates directly from get upon nitrates in the organic matter. hydrogen. that is. calcium. glycerin and fatty acids. of still other elements have been shown to be necessary also. and iron. food y as it is This reserve called.110 FUNDAMENTALS OF BIOLOGY They contain only carbon. Fats usually occur in liquid form in plants. as oils. and nitrogen. potassium.

carrying on their vital activities. a process that may go on in any living part of the plant. If a green plant is kept in darkness so that photosynthesis cannot go on. It has already been seen that roots and seeds are common storage organs. tubers. are always undergoing decomposition so that energy may be available. The commonest starch-digesting enzyme in plants is diastase. and so that they be eventually made into living matter. which in turn is dependent upon food manufacture. This is necessary in order is . fats. that they may that they utilize energy in the performance of their functions. Growth represents a predominance of the constructive phases of metabolism over the destructive ones. it represents the final stage in constructive metabolism. Plants and animals are machines in the sense Respiration. may pass by diffusion from cell to cell. and. This energy is derived from the breaking down of complex organic substances within their own living bodies. or bulbs) serve as organs of food accumulation.METABOLISM AND IRRITABILITY IN PLANTS 111 starch. and because they contain little water. Thus sugar itself and all the other kinds of foods made from sugar contribute to the actual organic substance of the plant. The conversion of starch to sugar at Digestion an example of a digestive change. of which there are many kinds. there is a steady loss of . If a paramecium is kept in water containing no food. Digestion in organisms is accomplished by the action of substances called enzymes. Unless renewed by the assimilation of additional food. All seeds contain reserve food. leaves. although often sugars. Reserve foods are mostly in an insoluble condition. and the process of digestion is merely a means of rendering foods soluble. either by the plant itself or by its offspring. as a consequence. while in many plants underground stems (rhizomes. Before reserve food can be utilized it must undergo digestion. and Assimilation. its dry weight decreases because its protoplasm is slowly being destroyed. the food is highly concentrated. some future time. Assimilation is the transformation of digested food into protoplasm. and ordinary stems Reserve food represents nourishment to be used is usually small. and proteins accumulate in varying amounts. we can actually observe a gradual decrease in size until death finally All organisms are constantly consuming energy in ensues. food substances. The amount of food stored in fleshy fruits. as well as living matter itself. Growth can take place only through assimilation.

Respiration is the means accomplished within the bodies of all plants and is essentially an oxidation process. the constructive phases of metabolism must counterbalance the destructive ones if the life of an organism is to be maintained. FUNDAMENTALS OF BIOLOGY Protoplasm must be built up because it is constantly being broken down. in the protoplasm. which is energy at rest. and through the roots plant. but in the daytime both Under favorable condiprocesses take place simultaneously. entering the plant through the stomata and lenticels. In other words. for. which goes on all the time. the hitter breaks down into simpler products with an accompanying release of energy for use by the living machine. Much misunderstanding often arises from failure to appreciate the significance of the gas exchanges that occur between green plants and the atmosphere. Oxygen is carried to all the living cells of the mainly through the intercellular spaces. this must be converted to kinetic energy. The chief products resulting from the decomposition of protoplasm through oxidation are water and carbon dioxide. . carbon dioxide is absorbed and oxygen is liberated. solar energy is absorbed when carbohyThis energy becomes stored drates are made by photosynthesis. as previously explained. At night only respiration is carried on. this is Respiration it is dissolved. which proceeds only in the daytime. To be utilized. of the oxygen is absorbed from the air (of which it forms about one-fifth). or energy in motion. The hitter passes out of the plant through the stomata and lenticels into the air. the gas exchange is just the reverse. The fundamental feature of respiration. and when food is assimilated it becomes incorporated All organic matter contains potential energy. Respiration goes on at all times in all living cells in all It involves the absorption of oxygen and plants and animals. and Most this is why all organisms need oxygen in order to live. is not this gas exchange but the destruction of organic matter within living cells in order that energy may be liberated for use in carrying on vital functions. in the food.112 weight. diffusing into the soil. Therefore it should be understood that in photosynthesis. however. the liberation of carbon dioxide. all vital The ultimate source of energy is the sunlight. When oxygen combines with protoplasm. but some also diffuses into the roots from the soil water in which by which animals. while in respiration.

Photosynthesis and respiration PHOTOSYNTHESIS Organic matter is may be contrasted as follows: RESPIRATION constructed Organic matter is destroyed and CO 2 are raw materials Oxygen is liberated Energy is stored 2 H O H^O and CO-> are waste products Oxygen is absorbed Energy is liberated Occurs in Occurs at all living tissues all Occurs only in green tissues Occurs only in daytime Irritability. the primary root grows downward. 80 and 81). plants react to than do slowly animals. the primary root always grows downward.METABOLISM AND IRRITABILITY IN PLANTS tions. there is much off than carbon dioxide. forcing its way through the soil. no matter in what position a seed is placed in the soil. 113 more oxygen given may however. however. are most Irritasensitive to stimuli. is placed in a horizontal position.The action It is a in seedlings. times In order that the various metabolic functions of the plant may be carried on most advantageously. light. while the stem pushes upward toward the light and air. Under ordinary bility in plants is conditions. itself is just as definite in common to all stimuli much more is Yet the response their cells are enclosed within rigid cell walls. when photosynthesis is active. the latter not even pass out of the plant but may be immediately utilized in the manufacture of sugar. which of responding to external influences. the root tip soon begins to turn downward again and the stem tip in the opposite direction (Figs. plants Young parts of plants. when it sprouts. and the mechanism of response is very since plants possess neither nerves nor muscles. of gravity can be seen to advantage well-known fact that. This is accomplished is chiefly through the property of irritability. it is necesvsary that all the vegetative organs be brought into the most favorable relations with their environment. and different. but older parts often respond as well. in as animals. the main stem grows upward. or stimuli. In fact. organisms. and moisture. If a Geotropism. such as root tips and elongating stems. is the power Irritability an inherent property of protoplasm and so In general. The chief stimulus involved seedling . manifested chiefly by the direction of growth and orientation that the various organs assume. The chief stimuli that call forth these responses are gravity. and the branches and leaves are held in a more or less horizontal position.

81. seedling of scarlet runner bean placed in a horizontal position The root tip is growing downward in response later. to the same stem respond stimulus. at right angles to the direction in which the stimulus is acting (Fig. and photographed 12 hours leaves and the stem.114 FUNDAMENTALS OF BIOLOGY It is evident that the root and is gravity. the stem away toward root manner. Older seedling of scarlet runner bean placed in a horizontal position later. 80. oriented in a horizontal plane. . Young and photographed 12 hours to the stimulus of gravity. the growing in these reactions FIG. 81). but in an exactly contrary the stimulus. FIG. Response to the stimulus of gravity is termed geotropism. that is. generally becoming it. Note the response on the part of both the from Leaves react to gravity also.

so that the stimulus of gravity acts in a new direction. the FIG. stimulated to grow faster. and in Fig. perpendicular to the light rays. The mechanism of response important to understand. Phototropism. it responds by a This is brought about by an unequal acceleration curvature. If a plant is placed in a horizontal position and slowly rotated so that the stimulus of gravity can act upon it with equal all sides. 82. called one of the most obvious expressions of irritability The response . 81. the upper side of the root. or by a twisting of the base of the blade lower side of the stem is itself. of growth on opposite sides of the organ Thus. A young mallow plant (Malva) which has been exposed to oneThe flat faces of the leaf blades are held sided illumination for several days. The leaf is oriented by a growth curvature of the petiole. of the shoot to light. intensity from is there is no reaction. 80. When an organ is placed in a different position from that in which it has been growing. in Fig. phototropism.METABOLISM AND IRRITABILITY IN PLANTS Mechanism of 115 is Response.

while the loaves bend so that the blades are perpendicular to the direction from which the is when a house plant coming (Fig. If a plant exposed to one-sided . illuminated from all sides but later from only one side. most plants. The plant was FKJ.) light rays. This is apparent placed near a window. the leaves assume a new position. (After first Noll. Note that the stem has bent toward the light and the loot away from it. S3. and that the light is of the latter is the stronger influence. as shown by direction of arrows. 45). With a change in the direction of the A mustard seedling growing with its root in water. This permits the absorption of a maximum amount of solar energy for use in photosynthesis. 82). stems grow upward.116 in FUNDAMENTALS OF HfOLOGY Under ordinary conditions of illumination. while the leaves have taken up a position at right angles to the light. the stem tips tending to grow toward the source of light. their orientation manifestly being such that the rays of light strike the flat surface of the blade at right angles (Fig. while most leaves assume a horizontal position. It is evident that the growth direction stem and the orientation of the leaves represent a definite response to the stimuli both of gravity and of light.

Although roots ordinarily do not react to the stimulus of light. its roots to grow from drier to more moist parts . there is no response to light. thus While this is not often apparent exhibiting hydrotropism. FIG. Although roots show little or no response to they are very sensitive to the stimulus of moisture. and in some plants. an experiment can readily be arranged in which the two stimuli act from different directions (Fig. under appropriate conditions. as the intensity of the stimulus is then equalized. tion. and the latter proves to be the stronger. Hydrotropism. 83).METABOLISM AND IRRITABILITY IN PLANTS illumination 117 is slowly rotated. because gravity and moisture commonly act from the same direclight. It is obviously an advantage to the plant for of the soil. Seedling of scarlet runner bean pinned to the underside of a sloping board covered with wet blotting paper. The stimuli of gravity and of moisture are acting from different directions. they grow away from its source (Fig. 84. Then it is seen that the influence of moisture is stronger than that of gravity. 84).

CHAPTER VIII THE LOWER ANIMAL GROUPS Although the simpler plants and animals resemble one another many ways. the gulf between plants and animals becomes more and more pronounced as we pass from the lower to the higher organisms. although in some cases this may be true.000 named and Like the plant kingdom. it is divided into a number of major groups or phyla. they have little in common. Each has reached a higher state of structural organization than those preceding it in the series. until finally. the members of each having The animal kingdom described species. it should be realized that there are exceptions to every one of them. in Cell walls present PLANTS and tissues ngid Cell walls absent ANIMALS and tissues internal soft Tissues less highly differentiated Organs external more highly Organs external and Tissues differentiated Chlorophyll usually present Food made by photosynthesis Response to stimuli slow Growth occurring throughout Stationary life Chlorophyll absent Food obtained from external sources Response to stimuli rapid Growth confined Motile to early life includes about 800. except for bade features. however. Each group should be regarded as standing for a different degree of progress from what was originally a more primitive condition. as given below. these differences. It must not be supposed. it will be well to have in mind the main differences between plants and animals as evidenced In considering especially by the more complex forms of life. These animal groups represent different stages in complexity and will be considered in an ascending sequence. Before beginning a study of the chief animal groups. that every group is related by descent to the one below it. bespeaking a common origin for all forms of life. the more complex members of each of the two organic kingdoms have become widely separated. certain fundamental structural features in common. In other words. 118 .

as in protozoans. but in some cases they are arranged in colonies. complexity in this group has come about through the specialization of parts of single cells. amoebic dysentery. and in the soil.e. but in some cases spore formation occurs. They are mostly marine in distribution. numbering over in size. conjugation may be temporary. Sponges are free swimming when young but later become permanently attached to objects in the water. in the bodies of other organisms. . or permanent. Ill and XIX. Statements of number of species in this and the next two chapters refer The actual number.000 species. are called metazoans. Fission is the prevailing method of reproduction among the protozoans. In addition to the types discussed in Chaps. constitute the lowest and unquestionably the oldest group in the animal kingdom. Animals that are not proto- zoans. Malaria. while some are parasites. is very much greater in some phyla. The individuals may be solitary but are usually group of 1 Sponges. of these simple creatures flourish in the ocean of the fossil record. Structural as in Paramecium. colonial. rather than by the formation of tissues. today and have since the beginning of the deep-sea oozes are Many made up chiefly of their shells. mention should be made of two large groups of marine protozoans whose cells secrete a shell around the protoplasm. comprising the simplest group of metazoans. all multicellular animals. in the ocean. but a few sponges occur in fresh water. Sexual reproduction is also common. as in Vorticella. and arthropods. and African sleeping sickness are human diseases caused by parasitic protozoans. All of them are unicellular and nearly all are microscopic The Protozoa. The" group numbers about 3. represent a transition between the protozoans and the other metazoans. In the one group (Foraminifera) the shell is composed Enormous numbers of lime.. i. They have had a long geologic history. Protozoans are found in fresh living water.000 species. as in the other groups. PORIFERA (SPONGES) The Porifera. which includes undiscovered species. being found in all seas. to described species. round worms. in the other (Radiolaria) of silica. As a rule the cells are solitary.THE LOWER ANIMAL GROUPS PROTOZOA 119 1 15. most probably. have arisen from an ancient protozoans.

ect. fl. a simple sponge (Sycon). after Shull. It has a large terminal opening known as the osculum. flagellum. then circulates through the canals. (A. collar cells X Grantia. . or branched. In all sponges the body by a great number of minute pores through It which water enters. cc. and finally leave's the body by way of the osculum. The body wall of many sponges is supported by 1 The outer and inner layers of a sponge seem not to correspond to the ectoderm and endoderm. but many are irregular. fan-shaped. 1 A. respectively. 85. ectoderm. separated This wall may is perforated be either simple or complex. spicule. but ordinarily it is folded in such a manner that a system of canals is formed. especially will cut it into two similar halves. . passes into the cloaca. B B. non-cellular mesoglea. sp.' ) The body wall of a sponge consists of only two layers of cells by a gelatinous. from Minchin. Most simple sponges are urn-shaped and radially symmetrical. A Fio. surrounding a central cavity called the cloaca (Fig. 85 A). Colonial colonial ones. 1 In the simplest sponges the body wall is not folded. portion of cross section of of ondoderm. Some form incrustations on rocks. "Principles of Animal Biology. the body has no definite symmetry. in Lankester's 2. sponges commonly are cup-like. portion of " Treatise on Zoology" . mes. of other metazoans. B. meaning that any plane which passes through the central axis of the body In many sponges.120 FUNDAMENTALS OF BIOLOGY The body of a simple sponge consists of a body wall Structure. since in the sponge's development the position of the primary layers apparently becomes reversed. mesoglea.

and transfer some of it to other They also engulf waste particles and pass outside the body with them. These may be loosely scattered or more or less interwoven to form a firm framework. 85). It is of interest to note that these muscle cells are stimulated directly. Sponges have simple tissues. often growing together to such an extent that the lines of separation between the individuals become obliterated. entering with the circulating water. Reproduction. protectively covered masses of cells that become detached from the colony in the late summer just before its death and produce new colonies the next spring. Budding is a common method of asexual reproduction in sponges. 85$). The fresh- water sponges form gemmules small. The cells sur- rounding the minute openings in the body wall are contractile close the pores in response to stimuli. A commercial or "bath sponge is merely the dried skeleton of a sponge colony of the latter type. Microscopic organisms and bits dead organic matter. in the canals.THE LOWER ANIMAL GROUPS 121 numerous small elements called spicules. no nerve cells being present. thus functioning as simple muscle cells. which line portions of the canals. In other is present consisting of a fibrous " organic material called spongin. are generally produced by each individual. but there is little cooperation sponges a supporting network among the cells in the performance of their functions. digest it. but since they ripen at different Fertilization takes place times. After ^the embryo has begome a small spherical . arising in the mesoglea. but each cell digests its own food. In the solitary sponges the buds become separated. and Water is drawn into the body and kept in circulation by means of ciliated collar cells. as in the protoDigested food may be transferred by diffusion from cell used as food. but in the colonial forms they remain attached. to cell. " The cloaca lining a canal engulfs and zoans. The amoeboid may take up food. Eggs and sperms. cells. Respiration and excretion take place directly between and the water in contact with them. are is not a digestive cavity. no definite organs are present. sponges are not self-fertilizing. wandering amoeboid of and gametes (Fig. Furthermore. Most of the cells in the body are of a simple type known as epithelium. which maybe composed of either lime or silica (Fig. In the mesoglea are spic ale-forming cells. the cells cells cells.

corals. Both are There are the two common . There are about 5. hydroids. terates. most of which are marine. External view of a hydra in expanded (A) and contracted (B) conditions.000 species.122 FUNDAMENTALS OF BIOLOGY of cells (a blastula). The hydra is one of the simplest of the coelenspecies: brown hydra (Hydra fusca) and the green hydra (H^ viridis). live in fresh water. Finally settles down COELENTERATA The coelenterates have many features in common with the sponges but show an advance in several important ways. include the hydras. it develops cilia mass and escapes into the it water as a free-swimming individual. Some of the coelenterates are solitary. but a few colonial. sea anemones. others are free swimming. etc. jellyfishes. and completes its development. 80. others attached. some are The Hydra. They B FIG.

and Van Ginn and Company. the body of the hydra is said This is a feature of all coelenterates.} inch. Cleave. hollow stalk attached at one end by a basal disk. 86). "General Zoology." (Redrawn from Linville. Kelly. Ovcrri/ FIG. The green hydra is somewhat smaller. 87. Locomotion may also .TtLK LOW UK ANIMAL UKUUt'ti live attached found in fresh-water ponds and streams where they to aquatic vegetation or to other objects in the water. Although attached. this difference being due to its power of contracting and expanding itself. after Parker. Because the parts extend outward in all directions from a common % to exhibit radial symmetry. Longitudinal section of Hydra. The body of a hydra consists of a cylindrical Structure. by permission. center like the spokes in a wheel. the animal can move from place to place by slowly gliding along on its basal disk. and bearing five to ten slender finger-like tentacles at its unattached end The length of the brown hydra varies from J^ to (Fig.

87). or gastrointernal termed the coelcnteron into a large cavity extends out fills the stalk but which not vascular cavity. The latter represents the primitive method characteristic of protozoans and sponges. The thread can be discharged into an animal coming in contact with the hydra. most expand and so function also as simple muscle cells. feeds chiefly on smaller aquatic animals. The The hydra endoderm also contains large digestive cells. and some" of it is passes by diffusion to the ectoderm. absorbs oxygen and gives off carbon dioxide and other waste products. can 87). thus carrying on respiration and excretion directly. being in contact with water. The food digested within the gastrovascular cavity is absorbed by the cells of the endoderm. Surrounding the central of two layers of cells. but these are not aggregated to form ganglia. . coming in contact with the tentacles. There is also present in both ectoderm and endoderm a loose network of simple nerve cells. The ectoderm contains peculiar stinging cells known as nematoWithin each cystSj which are most numerous on the tentacles. layers are composed mainly of of which have the ability to contract and epithelial cells. The digestive cells. are benumbed by the stinging cells and then conveyed by the tentacles to the mouth. Indigestible matter remains in the cavity. Nematocysts are used both for defence and for capturing prey. ways both inside the coelenteron and within individual cells. non- the mesoglea. the outer cavity is a body wall consisting the inner one the endoderm. the ectoderm and layer being called Between the ectoderm and endoderm cellular is a thin. into which digestive fluids are secreted by numerous small glandular cells. by thrusting out pseudopodia. engulf small solid particles of food. which in jellyfishes Both becomes very thick. releasing the basal disk. substance called nematocyst is a coiled hollow thread containing a poison. Digestion takes place within the coelentcron. gelatinous. only into the tentacles as well (Fig. and so digestion occurs in two. The tentacles surround a conical elevation called the hypoThis opens directly stome. Each "ell of the body. each of which has one or more cilia whose beating keeps the food in motion (Fig. in the center of which is the mouth. and taking hold with it in a new position.124 FUNDAMENTALS OF BIOLOGY be accomplished by bending over. being finally expelled through the mouth. which.

One marine hydroid. develop a set of tentacles. In the marine hydroids. and later become detached to form a new individual (Fig. 8. species. 88. after Parker and Haswell. colony. 10. a colony is formed. or spertnaries. After going into a short resting period. 1. Portion of a colony of discharged into the water as The free-swimming c e 1 1^ is knob-like a organ ovary formed in the ectoderm near It prothe base of the stalk. by permission. v . Sperms and eggs are borne on (Fig. may also reproduce by fission.) sheath. Sexual reproduction in Hydra is a simple process. mouth. 4. the new individuals produced by budding do not become separated. animals related to the hydra.THE LOWER ANIMAL GROUPS 125 Reproduction. the same or on different individuals. coelenteron. often consisting of thousands of indiThe hydra viduals (Fig. duces a single large egg. a great many sperms. which are . the coelenterates show a marked advance over the sponges. 87). called arise in depending on the The male organs. it then grows to the adult condition. As a result. but this method occurs rarely 154A). 6. The hydra reproduces both sexually and In asexual reproduction a bud may grow out from asexually. a 2. 87). stalk of of the sperms penetrates the ovary and fertilizes the egg. but remain permanently attached. 88). the stalk. Zoology. testes groups of two or three as conical elevations near the upper end of the stalk (Fig. enveloping reproductive branch. ectoderm. bud. Cambridge University Press. 9. 7. which becomes detached and gives rise to a free-swimming individual (medusa). 3. Obelia. " 11 (From Shipley and MacBride. In possessing definite sexual organs (testes and ovaries). endoderm. They are ectodermal in origin. but soon becomes free. 5. Each testis produces Fin. The zygote then gives rise to an embryo that undergoes part of its development within the ovary.

each of which bears a single polyp. Representative coelenterates. are enclosed in a chitinous sheath that also covers the stalk and is expanded into a cup at the base of each nutritive polyp. the medusa stage of a hydrozoan jellyfish (Poly orchis). natural size. branching forms attached to objects in the ocean (Fig. 89. the skeleton of a ooral. 88). A.126 FUNDAMENTALS OF BIOLOGY Other Coelenterates. B. lacking a mouth and tentacles. Most of the polyps are somewhat similar Fio. a sea anemone (Metridium). The coelenteron is continuous throughout the stalk and branches but .) to an individual hydra. after Emerton. (C. C. The latter. The hydroids are colonial. A colony consists of a stalk with numerous short branches. but some are differentiated as reproductive branches.

The body wall consists of but two layers of cells. Like the sponges. The sea anemones and corals are attached forms. Many oceanic islands have been built up by the gradual accumulation of their remains. the gametes being the only haploid cells in the life history (see p. the zygote giving rise to a polyp from which. but there are no systems. by thin A medusa stage is not In the corals a hard external framework of lime or of organic material is secreted by the individual animals in the colony. PLATYHELMINTHES (FLATWORMS) are animals constructed on a higher plan of organization than the coelenterates but have some primitive features in common with them. The presence of stinging cells is characteristic. These are cup-shaped individuals with a marginal row of tentacles and a central mouth. is divided into chambers present. A species living in fresh water. 257). and as parasites. and for this reason coelenterates are said to be A single cavity is present (the coelenteron) having but one opening (the mouth). a colony is developed. The flatworms common 1 fresh-water form is is Planaria (Fig. The life history of an asexual and a sexual 1 generation. This phenomenon also called . The medusae bear sexual organs. The tissues are simple but are more highly differentiated than in sponges.THE LOWER ANIMAL GROUPS 127 does not extend into the tentacles. since in the hydroids both generations are diploid.4). The reproductive branches produce buds that become detached from the colony and develop into free-swimming medusae (Fig. the former being solitary and the latter In both groups the individual is a colonial (Fig. Some have a polyp stage in the life history. 89. diploblastic.000 in the ocean. polyp in which the coelenteron partitions. The true jellyfishes resemble medusae but are larger and more alternate occurrence in the complex. 90). by budding. 89 B and C). They number about 6. A few simple organs are present. constituting the ectoderm and endoderm. the coelenterates are an ancient group and probably originated directly from protozoan ancestors. while two groups " alternation of generations" but is not the same as alternation of generations in the plant kingdom. Digestion occurs mainly in this cavity but also in the individual cells lining it. They are solitary and free swimming. of unlike form. Radial symmetry is a feature of all coelenterates. is known as metagenesis.

In the flatworms an in ectoderm \ FIG. cc-phala. extensible pharynx with an intestine that has three main branches. is differ- 90. passing through its longitudinal axis. of bilateral symmetry.128 of FUNDAMENTALS OF hlOLOGY that live upon certain of the higher and the tapeworms (Fig. their type of symmetry. Pharynx dorsiventrally. For example. becomes more marked in most of the higher groups. Planaria dorotofresh-water flat- a 10. but soon another layer. A bilaterally symmetrical animal has an (forward) and a posterior end. This feature continues subsequent groups.) Animals in which three primary cell layers are formed in the embryo are termed triploblastic. in all X (After Child. will divide the body into two approximately similar anterior (rear) halves. The sponges and diploblastic. one . known as cephalization. A primitive character retained by the flatworms is the presence of a single cavity (coelenteron) with just one opening (the mouth). worm. As are of their name implies. Intestine and a left and flat- a right side. is connected by a tubular. but nearly all the tapeworms flat worms parasitic of animals are the flukes members are colonial. Most the group are solitary. in the latter. and endoderm appear early development. entiated between them. called the mesoderm. 228). One of the most obvious differences between the coelenterates and the flat-worms is the presence. in Planaria the mouth. but only the two sides are alike. a dorsal (upper) and a ventral (lower) surface. This tendency. This means that only one Eye Nerve core/ plane. located near the middle of the ventral surface. the flat-worms exhibit a tendency to worms flattened Because organize a head region that physiologically dominates the rest of the body. but the cavity is usually highly branched. cells coelenterates are since only two layers of develop.

by burrowing through the soles of flesh of the 1 feet. and reproductive systems are present. this method is rare (Fig. more commonly. as in all of the higher groups. and mouse. Many roundworms are parasitic. and from it two lateral longitudinal body. It also attacks the pig. Although reproduction in the flatworms may occur by transverse fission. and 4. situated in the head region. rendering the head region Other nerves arise from the longitudinal particularly sensitive. in addition to more complex tissues and organs. Digestive/ nervous. The parasite lives in the intestine. and two A marked feature is exhibited by the flat worms in having. Digested food is carried the to all of the directly parts body by highly branched intestine. rat. or mass of nerve cells. . The excretory system consists of a network of tubes extending throughout the body. all three with many lateral extensions This (Fig. same individual but flatworms as a NEMATHELMINTHES (ROUNDWORMS) The round worms comprise over fresh and salt water. extend backward (Fig. nerve cords nerve cords and extend transversely. being transmitted from one to another when the an infected animal is eaten. entering the body either through the mouth or. in the soil.THE LOWER ANIMAL GROUPS anterior 129 posterior. 91). digestive tract corresponds to the gastro vascular of the cavity hydra. Man acquires the disease by eating improperly cooked pork containing encysted larvae. while respiration takes place directly through the surface. dog. muscular. other fibers are Muscle function than contraction. 1545). Tapeworms lack a digestive system.000 species occurring in in the bodies of other animals. but there is no circulatory or respiratory system. Almost all possess both testes and ovaries on the rule are not self -fertilizing. of the The body is mostly thread-like or cylindrical (Fig. 90). 90). Trichinclla causes a serious human disease known as trichinosis. Many small nerves extend forward from the ganglion. The hookworm (Necator) causes a human disease widespread throughout the southern part of the United States. definite systems. of the developed as sets of tissue's having no The nervous system of Planaria consists of a bilobcd ganglion. excretory. and in both cases? waste products of digestion are expelled through the mouth.

flatworms. in Yearbook U. familiar to every one. instead triploblastic animals with complex body systems. and an inner enteron. persist in all subsequent groups. Its length species of earthIt often studied. cavities two has to the exterior. appropriately designated as the segmented worms. S. the body surrounded by the body wall. the anus. Lumbricus terrestris is the one most burrows in rich moist soil.000 species of annelids occurring in fresh water. varies from about 6 to 12 inches. or digestive tube. are bilaterally symmetrical. Like . in the The earthworm is a familiar ocean. (From Cobb. Of the several common worms. land form. the sandworm a representative marine annelid. and a posterior one. The Earthworm. the enteron has two openings. and as parasites. General Features. greatly 1914. the mouth. The elongated cylindrical body of Lumbricus is pointed at either end and slightly flattened on the lower surface.) one. FIG. There are Coming now " about 6. an anterior A nematode worm (Xiphinema) that lives on the roots of plants. eatly magnified. These two features ANNELIDA (SEGMENTED WORMS) to the Annelida. with of having a single cavity *(coelenteron) just one opening an outer coelom. 91. Moreover. coming to the surface only at night or Its general appearance and habits are after a heavy rain. Department of Agriculture.130 FUNDAMENTALS OF BIOLOGY like Roundworms." we find a group that makes a great advance over those which have been thus far considered. while the leeches are well-known parasites. in soil. But.

It functions in reproduction. also seen in the roundworms^is carried on into all the higher animal groups. This important feature. As previously explained. first and last. bears four pairs of short They form four double rows The two and two ventral ones. anus. 93). . extends that a long digestive tube the anterior one being the mouth It has an opening at either end. is This divided the coelom. External view of the anterior portion of an earthworm as seen fioin the The thickened portion is the clitellum. In having two separate cavities coelom and enteron the body of the earthworm is built on the plan of a "tube within a tube" (Fig. intersecting the first above and beyond the mouth and backward on the dorsal surmetamere. large into compartments compartment and passing through the septa transverse partitions called septa. in last metamere. of earthworm consists of dead vegetable and animal matter often mixed with The food of the bits soil.THE LOWER ANIMAL GROUPS 131 the two lower groups of worms that have been considered. the ditellum. setae assist in locomotion by sticking into . The body wall of the earthworm is made up of several distinct It surlayers in which muscle tissues predominate (Fig. Digestive System. this means. very conspicuous. except the curved lateral bristles called setae. side. the annelids exhibit bilateral symmetry. a thickened band-like portion of the body wall comprising six or seven metameres. is FIG. each corresponding to a metamere. both externally and internally. or upper lip. Each metamere. which is thus not a complete Located about one-third of the way from the anterior end ring. earthworm consists of a linear series of ringlike segments called metameres or somites. 93). the entire length of the body. fluid-filled a rounds cavity. that only one plane. Within the coelom by is the enteron or alimentary canal. all of which are essenThe segmentation of the body is tially alike in form (Fig.the earth as the animal contracts. At the anterior end of the body the prostomium. 92). 92. slightly enlarged. passing through its longitudinal animal into two approximately similar halves. extends axis. will cut the The body of the face. being the the situated and the posterior one.

nephrost. n. worm. sub. longitudinal muscles. a narrow tube leading backward from the pharynx. circ. ?ieph. typhlosole. Soil passes through the alimentary canal. subneural vessel." dors. gland cells. a long straight tube constituting . cxt. n. anus. a muscular sac that draws food through In addition to the mouth and the mouth by i^^-*P^ coel veitt. v. co. 93. 94). mus. nephridium. (2) the esophagus. epid. (From Parker and HasweU. the earthworm actually eats ground. ventral blood vessel. after Marshall and Hurst. hep. setae. coelom dors. vent. follows: (1) the pharynx. cuticle. external opening of nephridium. long mus. set. nerve cord. cnteron. cut. nc. the organic matter in it is digested. circular muscle fibers.v out epid fteph hep extnepfa nephrost n. cod.) FIG. dorsal blood vessel. the digestive system comThese are as prises a number of specialized organs (Fig. a thin.walled enlargement of the digestive tube in which food is temporarily stored. "Textbook of Zoology" The Macmillan Company. vess. v. internal opening of nephridium. the gizzard.r set Cross section through the middle portion of the body of an earth epidermis. (3) the crop. generally its burrow. a thick-walled muscular organ where the food is (5) the intestine. ent. and the residue is deposited at the surface in the form of the familiar "castings. by . In way through the burrowing.132 FUNDAMENTALS OF KIOLOGY near the entrance to its It feeds at night. typh. (4) ground up.

A red pigment termed hemoof the globin is The blood dissolved in the plasma. and intestine (int). XV. pharynx muscular wall with thick (phw). ventral blood vessel (ubv). Excretory system includes Muscular ^system includes circular muscles (cm) and longinephridia (neph). These vessels extend . respectively. tudinal muscles (Im) Female reproductive sysventral nerve cord (nc). Circulatory system includes aortic arches (ht). and sperm ducts. the enteron. not shown. 93). overhung by the prostomium or upper lip (pr). is carried by a system of blood vessels and remains all inside these at times. The dorsal wall of the intestine is infolded so as to form a median ridge termed the typhlosole. Kelly. Following digestion. digestive glands (cgi to (ph) cfirs). erop(cr). "Princiides of Animal " Biology after Linville. Nervous system includes suprapharyngeal ganglion (br). the corpuscles being colorless. oviduct (ovd). and Van Cleave. side view. the dorsal and the ventral blood vessel. tris). dorsal blood vessel (dbv).) FIG. esophagus (oes). the soluble food passes by diffusion through the intestinal wall and is absorbed by the blood. and seminal receptacles (sri. the purpose of which is to increase the digestive arid absorptive surface (Fig. ST?). tem includes ovary (ov). gizzard (0i). In animals as complex as the earthworm. . which enclose testes.THE LOWER ANIMAL GROUPS 133 most of the digestive system. Digestive system includes the mouth (m). It is in the intestine that the Cells contained in the greatest part of digestion takes place. Male reproductive system includes seminal vesicles (svi to s#s). and other parts not shown. (From Shull. XX. inner lining of the intestinal wall secrete digestive enzymes into Others absorb the food after it has been digested. Circulatory System. the transfer of respiratory gases and of digested food is not possible without the aid of a special circulatory system. they are blood vessel called. Two principal vessels lie above and below the digestive tube. number of metamere. 94. The blood earthworm consists of a liquid plasma containing a large number of cells called corpuscles. subneural vessel (snv} and certain other vessels not here named. . V. not shown. X. Dissection of the anterior end of tho earthworm (Lumbricus tcrresI.

94). In addition to the blood. anterior end toward the right. external ports food to all parts of the possesses body. blood. the moist outer skin. Each nephridium has a ciliated funnel-like opening (nephrostome) in one metamere. 94 and 95). passes through the septum into the next metamere behind. Diagram to show the main structural features of a nephridium of the earthworm. 96. b. this movement being caused by pulsations set up both in the dorsal blood These principal vessels give rise vessel and in the aortic arches. (From Woodruff. the skin is kept moist so that this gas exchange can go on.) 1 Macmillan Company. down through the arches.' by permission. and then opens on the ventral surface of the body by means of a small pore (Figs. a. and backward through the ventral vessel. The earthworm no opening. beneath the ventral nerve cord and carries blood backward. takes up oxygen from the air and gives off carbon dioxide. containing small amoeboid cells. Each metamere. internal opening of nephridium.134 FUNDAMENTALS OF BIOLOGY body and are connected near the anterior end of five pairs of aortic arches that pass the length of the by means around the esophagus (Fig. By the secre- tion of mucus. a colorless fluid in the coelom. Excretion in the earthworm is also effected by the activity of the small amoeboid cells present in the . network of capillaries about the coiled glandular portion. These arise from the subneuml a long tube that lies vessel. The to many smaller branches that go to all parts of the body. has a pair of excretory organs called nephridia. The oxygen unites with the hemoglobin dissolved in the blood. These last one. carried by small vessels to The The Biology. "Foundations of special aerating organs. c. Blood flows forward through the dorsal vessel. and directly from the blood vessels by diffusion and convey them outside the body. also transFio. are long coiled tubes that collect metabolic waste products from the coelom. except the first three and Excretory System. blood from the various parts of the body is returned to the dorsal by means of a series of vessels encircling the body vessel posteriorly the parietal vessels.

ganglia of the ventral nerve cord with nerves emerging. . From the ganglia small nerves body wall and to the principal The largest ganglion is the internal organs. The female organs comprise a pair metamere. "Foundations of Biol- ogy" The Macmillan Company. a. An the first two. as the circumpharyngeal connectives. The former consist of two pairs of small testes located in the tenth and eleventh metameres and three pairs of large seminal vesicles. finally (From Woodruff. ducts with openings in the fifteenth metavesicles. 94). 96). a bilobed mass of nerve tissue situated in the third metamere pass to the d above the pharynx. Below the pharynx in the fourth metamere is a slightly just These smaller ganglion. A pair of tubular with external openings in the fourteenth metamere.) mere. double cord of nervous enlargement of this cord. System. by permission. suprapharyngeal ganglion. Reproductive hermaphroditic. first two ganglia are connected with each other by means of a pair of nerve cords. Diagram of dorsal view of the anterior portion of the nervous system of earthworm. The earthworm is This means that both male and female organs occur on the same individual (Fig. one of which passes around either side of the pharynx. Definite sense organs are lacking in the earthworm. circumphathe ryngeal c. The sperms are produced in FIG. glion. 6. sists The nervous system earthworm con- primarily of a ventral nerve cord extending the length of the body and located just beneath the ventral blood vessel (Fig. the subpharyngeal ganglion. being carried to the surface of the body by means of a pair of sperm. Nervous System. There are also two carries the eggs to the surface of the body. connectives. of small ovaries in the thirteenth oviducts. From it sensory nerves pass to the prostomium. subpharyngeal gan- the testes and stored in the seminal d. 135 These engulf solid waste particles and destroy of the them.THE LOWER ANIMAL GROUPS coelomic fluid. 96. but the skin is sensitive to touch known and to light. suprapharyngeal ganglion. called a ganglion. occurs in every metamere except It is really a fused tissue.

97. later. is hermaphroditic. Other Annelids. A similar stage appears in the development of certain other animal groups. sacs in earthworm are received arid stored. slips over the anterior end of the earthworm into the soil and forms a of the Most phorc. is At first the larva. carrying the Then fertilization occurs. The leeches and appendages. 97 A). swimming about by means of a circle of cilia. called a trocho- disk-shaped. A. (From Shipley and MacBride. and vice versa. a leech (Hirudo). The gelatinous tube. are more highly developed than the earthworm in that a head is differentiated from the rest of the body (Fig. upon hatching from the egg. after Oersted. eggs to the seminal receptacles.) fertilized eggs. Thus the eggs of one earthworm are fertilized by the sperms of the othef. and unsegmented. these passing from the seminal vesicles of the one worm to the seminal receptacles of the other. Later the clitellum secretes a gelatinous tube that is forced forward. is not self- Two ends pointing in come together with their anterior opposite directions and make an exchange of individuals sperms. cocoon. and other specialized organs. Some of the marine annelids. The setae may be in tuft-like masses borne on special lack setae locomotor appendages that also function as gills. tentacles. "Zoology" Cambridge University Press. A. within which the embryos develop. B. now containing B FIG. such as the sandwonn. marine annelids. as will be noted minute. Segmented worms. eyes. the sandwonn (Nereis). The head may bear jaws. by permission. Although the earthworm fertilizing. pass through a larval stage. a marine annelid. and have a sucker at either end of the .136 FUNDAMENTALS OF BIOLOGY which sperms from another it pairs of seminal receptacles.

and an enteron having both an anterior and a posterior opening. 98). 975). fishes. All the systems of the body show a higher degree of development than in the preceding groups. ray is Extending outward from this depression into each an ambulacral groove containing two or four rows of loco- motor organs. but as a rule all the metameres are alike and no body regions are present. show a great advance over the lower animal groups in being metameric. etc. but are never jointed. The lower. but most leeches are blood-sucking parasites. Embedded rays extending outward from a central disk (Fig. are spines present. sluggish in their habits. The body of a typical starfish is composed of five Structure. brittle stars. as their name implies. moving very slowly from place to place. longer but less numerous on the or oral than on the upper. mouth is situated in a depression in the center of the oral surface Numerous of the disk. of species of starfishes are found on Pacific coasts of North America. Q8A). a large coelom. . in the skin are calcareous plates forming a protective exoskeleton. ECHINODERMATA The cchinoderms are a peculiar group of exclusively marine animals numbering about 5. sea lilies. The Starfish. surface. The segmented worms. The anterior sucker contains leeches are aquatic. as well as They live on the sea bottom rather close to shore. the latter lying between the bases of two of the rays. small animals. Appendages are present in some Some of the annelids have a distinct cases.000 species. sea cucumbers. or aboral surface. They include the starsea urchins. head bearing sense organs. some living in fresh water and others A few of them live on dead organic matter or on in the ocean. 137 the mouth. Their outstanding characters are the presence of radial symmetry and a high degree of structural development as compared with Most of the echinoderms are motile but the preceding groups. A small red eyespot is present at the tip of each ray. A number both the Atlantic and along other seacoasts. On the aboral surface of the disk is a small central anus and a round perforated body known as the sieve plate. attacking larger animals. while the sea lilies are attached. the tube feet (Fig.THE LOWER ANIMAL GROUPS bcdy Most (Fig. They are triploblastic animals with bilateral symmetry.

98. A starfish common capitatus). aboral view. along the southern California coast A. B. natural size. oral view.138 FUNDAMENTALS OF BIOLOGY FIG. (Pisaater .

The is starfish possesses is but the coelom. and reproductive (Fig. Numerous short gills occur on the aboral surface of the body between the Definite organs of excretion are lacking. Ampullae Siomach pouch Anus Ovary Tube feet FIG. "General Zoology. nervous. 99). respiratory. with a colorless fluid containing amoeboid cells. with the anus by means of a very short intestine. and Van Cleave. The anus is not In each ray -are a pair of large greenish pyloric functional. filled no blood vessels. sends a pair of short pouches into each ray.THE LOWER ANIMAL GROUPS 139 The systems of the body show a high degree of development.) Dissection of the purple starfish (Asterias vulgaris).vascular. The alimentary canal is very short. as in the earthworm. The nervous system comprises a circular nerve cord in spines. the following being present: digestive. which large. 99. (After Linvttle. The large stomach. aboral view. water. . Oxygen and food are carried to the tissues by the coelomic fluid. by caeca (digestive glands). but the wandering amoeboid cells collect waste matter and pass outside the body with it. situated in the central It is connected disk. permission. Kelly. " Ginn and Company.

and enters the bulbous tube FIG. canal. radial tube feet. Reproduction. there are also cells lying in the skin. which are thereby exstone canal. feet. The starfish feeds largely on clams and oysters. water is forced back into the ampullae. . and the animal moves forward. adult condition. peculiarity of the starfish is the presence of a water-vascular system (Fig. important swimming. sieve plate." Henry pany. Water enters the sieve plate. while at the other end is a small ampulla. passes through the canal system. different individuals. 100). tube feet. Water. with a ring canal located in the central disk.vascular sys- t. ComHolt & of Zoology. 100. ceeds. and thus fertilization is external.) arc contracted. ampullae. worm-like ancestors. and to distal extends it the numerous attached. By contraction of the water is forced into the tem of the starfish: sp. wrapping its rays about the victim. tended and fastened to the sub(After Htrtwiff-Kingsley. Testes and ovaries in the starfish are of borne on sexual organs occurs at the pair base of each ray and during the breeding season occupies the Both the sperms and eggs are greater part of the coelom. and while ciliated but finally settles down and gradually passes into the This free-swimming larval stage. by means of a short tube called the stone canal. c. Loco. still in a very early stage of cilia and becomes free develops development (blastula stage) as development pronote to is It that. resembles the trochophore larva of certain marine annelids and suggests that the group may have been derived from bilaterally symmetrical. ring canal. by permission. characteristic of most echinoderms. ampullae. The released into the water. arid exerting a steady pull until it opens. fastening its tube feet to the shell. motion by this method is very slow. The sieve plate mentioned previously is con- A nected. r. small The tube feet are end of each tube foot forms a sucking disk. From into this a radial canal each ray. "Manual When the tube feetstratum. a bilaterally symmetrical larva is developed that remains A embryo develops at once. a.140 FUNDAMENTALS OF BIOLOGY many nerve the disk and a radial nerve cord in each ray.

X %. . a sea cucumber (Thyone). a sea urchin (Strongylocentrotus} X dmster). 5 . in lacking ambulacral grooves 101/1). and Locomotion is FIG. a brittle star (\ skeleton of sand dollar (D&n(/? X J^ <z/er Coe. 101. the animal swimming through the water. X %. fishes in 141 The brittle stars differ from the star- having slender rays . J>.sharply distinct from the disk (Fig.) accomplished not by the tube feet but by rapid movement of the The sea urchins rays. and sand dollars are covered with spines but do not have . A 5- y B. Representative echinoderms.THE LOWER ANIMAL Other Echinoderms.

The mouth. The echinoderms cal are an aberrant group of radially symmetri- animals probably derived from worm-like ancestors but not in the direct line of descent of any of the higher groups. lacking cephalized. The group is characterized three special features. The sea urchins are globular and There are no ambulacral grooves. confined to five double rows. record extending far back were particularly abundant . or may be absent. The crinoids. In of centralization such a as bodily activities. during certain ancient times. with five branched tentacles covered with tube feet. FUNDAMENTALS OF BIOLOGY 1015 arid C). but tube feet arc present and borne by five pairs of radiating calcareous plates. between which lie five other pairs of plates. and are peculiar in being attached by means of a long stalk. (3) by a water-vascular system. They are all triploblastic animals with a well developed coelom. The tube feet may be tentacles that are used in feeding. having small calcareous plates but no spines (Fig. viz. (1) a spiny calcareous skin. are mostly deep-sea animals. scattered over the whole surface of the body.142 free rays (Fig. bilaterally symmetrical animals possess. is surrounded by five branched the sand dollars flat. or sea lilies. having two openings. the echinoderms have emphasized an in i progressive type of organization. 101D). In the sea cucumbers the skin is soft and leathery. The echinoderms have left a fossil lilies The sea into geologic history. the enteron. in most cases. (2) the occurrence of the organs in fives or in multiples of five. situated at one end of the elongated body..

where they lie partly buried in the sand or mud. these are inexact terms but convenient ones to use They are inexact because the groups to which they refer cannot be sharply set off from each other. and lakes. of unsegmented body being encased in a calcareous shell composed two lateral valves (Fig. form the subject of the next chapter. backbone. cuttlefishes. slugs. and on land. around which may be seen the concentric lines of The umbo is the oldest part of the valve. being represented by such forms as mussels. are common mollusks closely resembling the clams. 143 . in the ocean. ponds.CHAPTER IX THE HIGHER INVERTEBRATES The seven animal groups presented in the last chapter. clams. animal kingdom. the soft. oysters. The fresh-water mussels. External Features. Mussels are bivalve mollusks. and octopuses. invertebrates those without Like flowering and non-flowering plants. Each valve bears a small rounded elevation called dorsal edges the umbo. 102). squids. They inhabit the bottoms of streams. constituting the "higher invertebrates are " the while the in the highest group vertebrates/ discussed. in being bilaterally symmetrical many They and more highly developed in include about 72. The Fresh-water Mussel. all of which are marine.000 species living in fresh water. may be conveniently In this chapter two " additional groups. together with several others of minor importance. Projecting growth. snails. MOLLUSCA of animals with a Like the echinoderms. the mollusks are a non-metameric group number of special features but differ from them ways. These are united along their by a-n elastic hinge ligament that keeps the ventral edges open. Vertereferred to as the "lower invertebrates/' ' brates are animals with a one. of which hundreds of species are known.

. one located at FIG. 103). and the space that it encloses is the mantle cavity. Passing over the gills. and Van Cleave. the contraction of the two thick adductor muscles. enters the mantle cavity through the ventral or incurrcnt siphon and leaves through the dorsal or Mantle Cavity. one-half natural size.) either mantle cavity. Ihe valves may be drawn together. the organ of locomotion (Fig. They are covered with cilia whose beating draws in water through the ventral siphon. as well as bits of dead organic matter. (From Linville. which sac thin muscular material composing it is secreted by a The lies just inside it and encloses the internal organs. bringing food and oxygen into the body and pair of large plate-like removing waste materials from it. The sac is the By mantle. The food of the mussel consists of microscopic plants and Food is guided animals. Water excurrent siphon. It permits the mussel to plow its way slowly through the sand or mud. (/ills occurs along the right and left sides of the mantle cavity A (Fig. 102). "General Zoology. The calcareous shell consists of three distinct layers. Kdly. by permission. the water is deprived of its dissolved oxygen. At the posterior end of the body is a pair of short tubular siphons end of the (Fig. Living fresh-water mussel (Unio complanatus)." Ginn and Company.144 FUNDAMENTALS OF BIOLOGY vcntrally from the space between the valves and directed downward is a large muscular foot. 102). 102.

(Modified after Jewell model./. 103. rectum. int. narrow. aur. stomach. cerebral ganglia. anus. kidney. anterior artery. mouth. which is very small. shell. pa. coiled intestine. aa. Surrounding the stomach is a large greenish digestive gland. dg. mo. Ip. Leading from the stomach is a long. intestine. gills Blood coming from the and mantle folds. cona median thick. ventral siphon. ventricle. m. foot. digestive gland. labial palps. auricle. ur.) communicates with it by ducts. gills. The mouth. ureter. which finally passes through the The rectum heart. dorsal siphon. visceral ganglia. vs. mantle: s. adductor muscle. is located between the anterior adductor muscle and the base of the foot (Fig. The sists of heart. enclosed by a delicate sac (the pericardium). extends from the heart to the anus. pg. pcj mt ov rn g s vq vs f fp FIG. posterior artery. st. ven. pericardium.THE HIGHER INVERTEBRATES to the 145 mouth by two pairs of labial palps that surround it. ov. ovary.walled ventricle and two lateral auricles. 103). . siphon. ad. pedal ganglia. It is connected with the stomach by means of a short esophagus. the latter being located above It communicates with the dorsal the posterior adductor muscle. without of course communicating with it. es. p. where it is aerated. which ad k ad ds mo. vg. an. g. r. esophagus. eg. ds. Dissection of fresh-water mussel. k.

The blood now makes its way to a large vein that goes to the Thence it kidneys. Waste The products are carried through a pair of short tubular ureters into the mantle cavity. Reproduction and Development. situated at the base of the posterior adductor muscle (Fig. These give off many small branches with open ends from which the blood passes into a system of spaces called sinuses. to the gills. The sperms leave the body of the male through the dorsal siphon and enter the ventral siphon of the female with the incoming water. finally passing out of the body through the dorsal siphon. The nervous system of the mussel is very It consists primitive. attach themselves to the After leaving the mother. 103). This the blood forward through the anterior artery contracts. (2) the pedal ganglia. The mussel has no organs of special sense. formed mussel (Fig. (3) the visceral ganglia. a feature correlated with its sluggish life. Nervous System. spongy kidneys situated just beneath the pericardium and communicating with it by means of a short duct. Finally the blood returns to the heart through small The blood of mollusks consists of innumerable colorless is corpuscles suspended in a liquid plasma in which bluish pigment called hemocyanin. The sperms and eggs pass through a pair of ducts into the mantle cavity. oxygen. the glochidia The young gills or fins of fishes. but the edges of the mantle folds and the siphons are sensitive to stimuli. such as eyes or tentacles. Nerve cords connect the pedal ganglia with the others. vessels.146 first FUNDAMENTALS OF enters the auricles BIOLOCfY and then passes into the ventricle. life now leads a parasitic as development proceeds. the sexes of freshwater mussels are separate. consist of a pair of cream-colored masses lying within the coils of the intestine. finally . The tcstes and ovaries^ which are very similar in appearance. these are divisions of the coelom. located in the foot. fertilization gill region. of the cerebral three of ganglia: (1) pairs ganglia. where it gives up its nitrogenous wastes. As a rule. it carbon dioxide and takes up liberates where goes. lying just behind muscle adductor and with each other by connected anterior the cord slender nerve of a over the means passing esophagus. dissolved a excretory organs of the miLssel comprise a pair of dark brown. forcing and backward through the posterior artery. and early embryonic development occurring in the Soon a peculiar larva known as a glochidium is 104).

cuttlefishes. The mantle folds bear numerous slender tentacles and a row of eyes. being asymmetrical owing to the necesThe chambered sity of accommodating it within a spiral shell. About 40 tentacles form a circle around the mouth. a rare deep-sea animal of the South Pacific Ocean. The or for capturing prey mantle forms a tubular sac that encloses the swimming body except the head. 105J2). None of the bivalve mollusks has a head. swim through the Scallops water by rapidly opening and closing their Shell FIG. the squids and cuttlefishes a entire Two large eyes are present. ductor muscle. become obscured most of the body. Here the foot is divided into either 8 or 10 tentacles. which is genAll of them have a erally encased in a hard calcareous shell. a ciliated free- swimming larva is developed similar to that seen in the annelids and echinoderms. each representing a space in which the animal formerly lived. a Hoof: Thread hatchet-shaped foot adapted for and a pair of siphons. nautilus. In pair of lateral folds of the mantle forms which assist in swimming. (glo- chidium). The bilateral symmetry that characterizes other mollusks has in the case of the snails and their relatives. Clams are marine forms resembling mussels in having a bivalve shell. 105/1). is present. much (After Bal- valves. Muscle Larva of mussel enlarged. fins. such. their two valves being unequal in size. All mollusks . as oysters. have a soft non-metameric body. broad and flat and is used for crawling. or internal and reduced to a small flat plate. bearing eyes arid tentacles. In many mollusks. The snails and their relatives usually have sisting of a spiral shell conshell is only one valve (Fig. The foot A is distinct head. and Oysters scallops are marine bivalves that lack a foot and siphons and have only one addigging. the shell is either absent. Other Mollusks. but in some cases a lacking. In the related squids. has a coiled shell divided into compartments.THE HIGHER INVERTEBRATES 147 leaving the fish and dropping to the bottom to assume an independent life.) 104. except the head and foot. and octopuses. Oysters lie flat on the sea bottom. which are used in (Fig. four.

A.. a small ortopus (Polypus). but characteristic organ. A distinct head may or may not metrical and triploblastic. one-half natural size. . and in Mollusks are bilaterally symmost cases a mantle present. the fact that the larvae of many mollusks are ciliated and free swimming suggests a remote affinity with the annelids and ecliinoderms. natural size. the foot. Although their relationships to other groups is not obvious. 105. a land snail (Helix). is well The digestive system occur.-Kepresentati vo molluskg. B. developed and has both the coelom is small.148 FUNDAMENTALS OF BIOLOGY which is variously modified. is FIG. Mollusks make their mouth and anus. appearance as fossils in very ancient rocks and have remained abundant throughout geologic history.

they are Antenna An-f-ennu/e f?os I rum Cepha/ofhorax 'Abdomen Uropod . one-half closely related to the segmented worms. The chief classes are .THE HIGHER INVERTEBRATES ARTHROPODA 149 Numerically the arthropods surpass all the other animal groups combined. notably in the possession of jointed appendages." The lowest class are almost exclusively aquatic. there being over 640. The number arthropods are more highly developed than the annelids in a of important respects. Telson view of western crayfish natural size. there are about four times as many arthropods as all the other known species of animals taken together! Although having few features in common with the echinoderms and mollusks. showing striking evidence of having been derived from them in the distant past.000 species. the name arthropod meaning "jointed foot. In other words.Flu. but nearly all the other members of the group live on land. Dorsal (Astacus trowbridyii) . 100.

The cervical groove. Each eye is composed of about 2. called the carapace. is present on all parts of the body except the ventral surface of the abdomen. the head of five.. a basal portion (protopodite) giving rise to an inner branch (endoIn the embryo all podite) and an outer branch (exopodite). but later they become modified in accordance with the functions that they are to perform.500 facets (simple eyes). thus permitting movement to take place. One of the swmmerets from the third. are fresh-water crustaceans very similar to the lobsters. the rostrum. arid arachnids. marks the line of union between the head and thorax. is made up of metameres. fourth. on either side of which is a large stalked compound eye at the end of a movable stalk. The exoskeleton is shed periodically so that can distinct plate. The Appendages. Some kinds live in moist meadows where they make burrows in larger and live in the sea. covers occur. Frequently the exopodite is lacking. including a number of species. all but the last one bear a pair of jointed appendages. be considered separately. Each of these eight. myriapods. a depression across the extending carapace. The most significant fact about them is that. thorax. A firm outer covering com- posed of chitinj a horny organic material. They crawl on the ground. growth the top and sides of the cephalothorax. its presence obscuring the A in this region. viz. This exoskeleton not only protects the internal organs but serves as a place for the attachment of muscles. and abdomen. . which are the bottom of ponds and streams. The crayfishes. External Features. in spite of their diversity. and impregnated with lime. which form separate images. the crustaceans. The body of the crayfish is composed of a head. These will The Crayfish. the cephalothorax (Fig. 106).150 FUNDAMENTALS OF BIOLOGY insects. Of the 20 metameres in the body of the crayfish. Anteriorly the carapace is prolonged into a metamerism hard beak. the thorax of and the abdomen of seven. they are all constructed according to the same fundamental pattern and so are said to be homologous. concealing themselves beneath stones and logs. the appendages are two-branched and otherwise similar. or fifth abdominal segment shows the structural plan common to all the appendages. It is thin and flexible where joints occur. but the first two regions are fused to form a single division.

or last metamere. and their functions are given in the above table. their location. esophagus. and a pair of The circulatory system comlarge digestive glands (Fig. together with the telson. fan-shaped tail effectively used in swimming backward. The names of the appendages. anus. form a flat. composed function is of a plasma containing to transport both food numerous amoeboid cells. The appendages Internal Anatomy.THE HIGHER INVERTEBRATES 151 of the nineteenth metamere (the uropods). 107). stomach. intestine. as in the walking legs. sists of The digestive system of the crayfish con- a mouth. its and oxygen to the tissues and . The blood is prises principally a heart and seven main arteries.

external opening of sperm duct. entering through three pairs of lateral openings. ar. gills. as in The blood leaves the heart through the arteries. anus. chelipod. u. cephalothorax.152 FUNDAMENTALS OF BIOLOGY The crayfish to remove metabolic waste products from them. constitute the respiratory organs. . "General Zoology. stomach. spg. c. testis. r. intestine. and six in the abdomen. ms. 107. has colorless blood. subesophageal ganglion. it returns to the heart. (After Linville. Dissection of male crayfish (Cambarus limosus]. located beneath the carapace on each side of the cephalothorax. antenna. flows into smaller vessels with open ends.} XL or three rows of plume-like gills. situated in the ventral part of the head. sbg. al. an. sw. uropod. compound eye. and a pair of green glands. where the nerve cord is Nerves pass double. by permission. artery. where it is single. general construction of the nervous system is A ventral nerve cord extends the length of the body beneath the alimentary canal. sd. supra-esophageal ganglion. h. a. ch. s. dg. The similar to that of the earthworm. after being aerated. The blood now goes to a large ventral sinus and then by veins to the from which. third maxilliped. sw FI<J. ct. abdomen. t. nerve cord.s or spaces that surround the internal organs. cs. nc. digestive gland. i. esophagus. Their external openings occur on the basal segments of the Two antennae. antennule. Seven ganglia occur in the cephalothorax. swimmeret. and from these passes out into swu. e." Ginn and Company. rostrum. but that of most crustaceans is blue. and Van Cleave. heart. comprises the organs of excretion. the mollusks.

numbering about 18. They have a calcareous shell consisting of several pieces. cephalothorax. (Fig. crabs. the annulus. shrimps. lobsters. ural size. have fused together. Other Crustaceans. partial Most crustaceans live in the ocean. 108). and as adults show little resemblance to other crustaceans. as the case may be. to various objects in the sea. . The sperms are deposited in a shallow cup. Striped shore crab (Pachygrapsus crassijjc s) three-quarters natAs in other crabs. is connected by a pair of slender cords with the second one. young remain attached to the proceeds. There is only one testis or ovary present. posts. by means of of these represents several ganglia that Each Reproduction. the abdomen is permanently folded under tho . lying above the esophagus.000 species and including the crayfishes. female between the fourth pair of walking legs. located beneath the esophagus. and ship bottoms.. and here they The eggs are fertilized when laid are stored for a long time. The a pair of ducts. He. sow bugs. swimmerets until able to care for themselves. some and a few on land. the swimmerets as development to and then become fastened After the hatching. but each is connected. such as rocks. which occurs in the >-> FIG. prawns. fresh water. Some have adopted a life in of The barnacles live attached or complete parasitism. 108. barnacles. sexual organs of the crayfish are always borne on separate individuals. to a pore situated at the base of each of the fourth walking legs in the male. The crustaceans arc mainly an aquatic group. the second in the female.THE HIGHER INVERTEBRATES from all 153 the ganglia to various parts of the body. The first ganglion.

Except ages. and each of the head (except the last two) bears a single .154 FUNDAMENTALS OF BIOLOGY Crabs are peculiar in having the abdomen permanently folded under the cephalothorax. The myriapods are a terrestrial group of comprising the millipedes and centipedes These forms are more worm-like than the three (Fig. 247). 109). B. other classes of arthropods. the respiratory organs consist of a pair of gills. 109. in the lowest forms. The Myriapods. about 2. eyes. into which the body of a crustacean is divided. Millipedes are mostly cylindrical every metamere back of the head (except the first one) bearing two pairs of short jointed legs. metamere back Centipedes are flattened dorsiventrally. ancient and most primitive arthropods were the a group of crustaceans (Fig. having a head but no differentiation between thorax and abdomen. and typically each one bears a pair of branched. The head. . which is commonly broader than long. A.000 species. jointed appendThe exoskeleton contains both chitin and lime. matter. a millipede (Spirobolus) natural size. . and the excretory organs of a pair of green glands. In most cases the head bears two pairs of antennae and a pair of compound The number of metameres is rarely more than twenty. or the head and thorax may be fused to form a cephalothorax. but have been extinct The most for many millions of years. These forms were very abundant during early geologic times. a centipede (Scolope ndra) slightly reduced. FIG. and abdomen. may be entirely separate. thorax. trilobites. Myriapods. They are sluggish in their habits and most of them feed entirely upon vegetable in shape.

head. ant. The head of the grasshopper is composed of five or six mctameres. which have become fused to such an extent that their individuality is lost. The grasshoppers. and. spiracle. It is encased in a hard chitinous covering (exo skeleton). /to. t?. h.THE HIGHER INVERTEBRATES pair of 155 Centipedes are very active creatures. They live in grassy places and feed upon vegetafw he ant rn t3 ti U +a li 3 sp External anatomy of male grasshopper. hw. l\. and most of them have two simple eyes. External Features. abd. compound eye. tn. thorax. antenna. hind (After Packard. The Grasshopper. Both millipedes and centipedes have only one pair of antennae. 110. s. a thorax. each consisting of a small group of simple eyes that are not so closely organized as the facets of a compound eye. three of them making up the thorax and ten the abdomen. insects. fore wing. are widely distributed insects comprising a large number of species. or ocelli. legs. 1 Their respiratory and jointed legs. abdomen. In some localities they are very abundant during the late of summer and autumn. any one of which will serve to illustrate the general features of insect morphology. fa. the latter being arranged in the form of a triangle 1 Some of the millipedes have a pair of aggregate eyes. which is shed periodically to permit growth to occur. J FIG. X<3. . 110). e. tion. or locusts.} wing. The head bears a pair of large compound eyes and three simple eyes. and an abdomen (Fig. worms. auditory organ. 1. The body an insect consists of three distinct regions: a head. and other small animals. Is. The other metameres are distinct and easily recognized. capturing excretory organs are similar to those of insects.

one pair on each metarnere. which In the grassis incomplete. arc united laterally to form the labium (lower bles The mandi- of maxillae. narrow. although perhaps only n. and but slightly hardened.156 FUNDAMENTALS OF BIOLOGY of the head. and second maxillae 1 The second lit)). which forms its own image. maxillae. . The thorax bears three pairs of jointed legs. memof the body wall. mouth parts such as charac- Mouth X 4. they are the antennae ing jaws). together with the flap-like labrum (upper lip) and the small hypo- and two pairs pharynx (tongue). first maxillae. hopper. which are attached to the second and third thoracic metameres. but in all except the very lowest insects no abdominal appendages are present. lab. hopper the first abdominal metamere. terize generalized The mouth ably parts of insects that obtain their food by sucking are consider- modified. on the front is As in the crayfish. second maxillae forming labium. and folded beneath the fore wings like a fan when the insect is not flying. mx. 111). . Wings are not appendages but merely outgrowths Insects differ considerably in the character of their wings. each compound eye composed each of thousand hexagonal units called facets. of several grasshopper hears four pairs of In the order jointed appendages. branaceous. constitute the mouth parts of the grasshopper (Fig. Ibr. hypopharynx. 111. parts of grasshihruin md. mandible. first mandibles (bit- maxillae. They are typical bitinsects. and are adapted for jumping. of their occurrence. although con- structed according to the same general plan as those of the grasshopper. The head of the partial one. In the more primitive arthropods all or nearly all the abdominal metameres bear appendages. while the hind wings are large. ing FIG. in the grasshopper the fore wings are thin. Almost all insects have two pairs of wings. hyp. Three pairs of legs are characteristic of practiIn the grasshopper the hind legs are very large cally all insects. (feelers).

Circulatory abdominal wall that permit all Situated directly below the dorsal System. The last abdominal metamere in the female is modified to form an egg-laying organ. anus. occur in pairs on all the metameres Spiracles. s. 112). consists of a mouth. g. a large thin-walled stomach. a long slightly coiled intestine. and six or eight large gastric caeca opening into the anterior end of the stomach. mt. (Modified after Jewell model. esophagus. nerve cord. Certain digestive glands are also present.. gastric caeca. antenna. and c ant he ov ^^ int an m sbg s ml in-f . c. while in the male two abdominal ones. an. is a long. heart. 112. or breathing pores. stomach. It is partially divided into eight chambers and furnished with valves the blood to flow forward. gc. oviduct. thick-walled. nc. the oviposiit has two structures for transferring sperms. mouth. a small spg es muscular gizzard. m.THE HIGHER INVERTEBRATES 157 bears a pair of auditory organs.} FIG. intestine. Malpighian tubes. gizzard. a large sac-like crop. crop. sg. ovd. contractile heart. From it a slender branched vessel extends into the head. spg. ovary. viz. h. These secrete digestive fluids into the alimentary canal. es. subesophageal ganglion. As in the mollusks and . ovd Dissection of female grasshopper (M elanoplus) ant. supra?sophageal ganglion. int. salivary glands. several pairs of tubular salivary glands emptying into the mouth. which ends at the anus (Fig. back of the head except the third thoracic segment and the last tor. a short curved esophagus. The digestive system of the grasshopper Digestive System. ov. sbg.

158 FUNDAMENTALS OF BIOLOGY in the other arthropods. respectively. tubes. A pair of testes. the esophagus. and mouth parts. entering through lateral openings. air are caused by rhythmic contractions of the body wall. consisting of a so well developed as that of the crayfish. are borne on separate individuals. ganglia. esophageal ganglion and the subesophageal ganglion. has retained paired metameric nephridia such as characterize the segmented their thin walls. oxygen or carbon dioxide. Nervous System. In all insects male and female organs Reproductive System. only of the For this reason the circulatory system grasshopper is not colorless. Respiratory System. the two parts of which It sends are separate in the thorax and united in the abdomen. antennae. The excretory organs of the grasshopper Excretory System. 145). consisting . the blood of insects is plasma and numerous is that it carries no feature a amoeboid corpuscles. They cords are connected with each other encircling by means of a pair of nerve Nerves extend from these Of the other ganglia to the eyes. one above and the ganglia arc present in the head. which form an internal network carrying oxygen directly to the tissues in all parts of the body (Fig. These are throughout the intestine. out nerves to various parts of the body. tory system and tubes called tracheae. which give buoyancy to the insect in flight. The tracheary system is connected with large air sacs. It is general features of the nervous system Two large are similar to those of the earthworm and crayfish. The These are called the suprasmaller one below the esophagus. all connected by a double ventral nerve cord. consist delicate. which It finally returns to the heart. of the end coelom and opening into the anterior They remove nitrogenous wastes from the blood by absorption through highly coiled structures extending an interesting fact that a very primitive arthropod (1 \ripatus). the blood flows out into sinuses. it communicates with the entirely distinct from the circulaconsists of a large number of fine connected This is The entrance and exit of outside air through the spiracles. but peculiar food and nitrogenous wastes. of a large number of Malpighian. As in the crayfish. three occur in the thorax and five in the abdomen. Many of the ganglia have been formed by the fusion of smaller ones. the larger worms. belonging to a class of its own. are divisions of the coclora.

dragonfly D chinch bug (Blissus). 113. potato beetle (Leptinotarsa) . C\ cabbage butterfly (Pieris).} (Libdlula). by permission.THE HIGHER INVERTEHKATKS 159 A. t FIG. . F. katydid (Microcentrum) . after various authors. "Outlines of General Zoology" The Macmillan Company. E. -Representative insects. house fly (Redrawn from Newman. (Musca). B.

140). and cockroaches have biting mouth Their anterior wings are straight and leathery. Sucking mouth parts are present (Fig. wingless condition. last located above the intestine. Except in a few primitive forms. The insects.000 species. but both pairs of parts. the adults except for their small size. larvae worm-like the young being (see pp. the posterior The wings are membranaceous and about equal in size (Fig. 241-242). The most primitive insects are the small wingless creatures without springtails and fish moths The compound eyes and in some cases with abdominal appendages. crickets. 1135). The flies and mosquitoes also have sucking mouth parts but have only one The beetles have biting mouth parts pair of wings (Fig. ants. is connected with the duct. 113D). In the and wasps all the wings are membranaceous. the hind pair being smaller than the fore pair. abdominal appendages are absent. but in some the fore wings are thickened at the base. The excretory system is composed of Malpighian tubes. and include an enormous variety of forms. One pair of antennae is present. which occurs within the female's body. The two The ovaries the eggs reach the ovipositor through a pair of After fertilization. the latter consisting of tracheae. 113A). 113F). numbering over 600. bodily proportions. The circulatory and respiratory systems are entirely distinct from thorax. arid undergo a metamorphosis. bees. each other. . each being metamere by means of a slender sperm are also tubular and situated in abdomen. Nearly all insects have six legs and two pairs of wings. covering membranaceous wings when folded (Fig. In all insects the body has three separate divisions: head. Other Insects. the eggs arc laid in the ground in masses of 20 to 35 and These are essentially like later hatch into young called nymphs. and most cases there are two compound and three simple eyes. oviducts. Most of the true bugs have membranaceous wings. 113JS).160 of a FUNDAMENTALS OF BIOLOGY mass of tubules. and hard fore wings that meet in a straight line and serve as covers for the membranaceous hind wings (Fig. in and abdomen. The dragonflics also have biting mouth parts. higher insects constitute the largest group of arthropods. 113C). butterflies and moths have sucking mouth parts and scaly wings (Fig. The mouth parts are adapted for biting and sucking (Fig. The grasshoppers.

Their abdomen is differentiated into two portions and bears a terminal sting. Arachnid*. a ful. The is externally indistinct in the other arachnids. Here belong the spiders. rus hirsutus) . except in the are without abdominal appendages. maxillae that are often and claw-like have power large scorpions arthropods numbering about 20. Wings are never Arachnids have four of developed.THE HIGHER INVERTEBRATES The Arachnids. 114). but longbgs. In nearly all arachnids the body consists of a cephalothorax and an abdomen. t$. natural size. mites.000 Fiu. daddy longlegs. rarely The excretory organs consist of Malpighian tubes. The arachnids 161 constitute a distinct class of species. pairs legs. A garden spider (Aryiopv aurantia). but of simple eyes are present. Antennae and compound eyes are lacking. tion takes place in characteristic organs called lung books. and. entirely a variable number in tracheae. king crabs. and ticks Metamerism is apparent in the scorpions and daddy (Fig. 11-1. . scorpions. Respiraking crabs.

162 FUNDAMENTALS OF BIOLOGY General Features of Arthropods. is In the crustaceans the body covering impregnated with lime. Insects have a distinct head. triploblastic. 'Two pairs in tire millipedes. They differ in having a reduced coclom. distinct body regions. and abdomen. thorax. . a chitinous outer covering. but with no distinction between thorax and abdomen. In the two primitive classes (crustaceans and myriapods) nearly every segment. as a rule. metameric animals with highly developed systems and an to the enteron with two external openings. but in the more advanced groups (insects and arachnids) appendages have disappeared from the abdomen. The arthropods are similar segmented worms in being bilaterally symmetrical. bears a pair 1 of jointed appendages. while in the arachnids and most crustaceans the head and thorax are fused. and jointed appendages. The myriapods have retained a worm-like body with a head.

but having certain fundamental features in common with tho backboned animals. at least in early life. In the vertebrates. are intermediate. which is then purely an embryonic structure. and in the embryo stage of development. The chordates arc characterized by three distinctive features. this structure may persist throughout life or may later disappear. triploblastic animals with a coelom and an onteron. invertebrates. CHORDATA The chordates.CHAPTER X THE VERTEBRATES or absence of a backbone forms the basis of vertebrates and invertebrates. jointed appendages are present. a segmental arrangement of certain parts is apparent. but. however. a vertebral column (or "backbone") arises which is made up of a linear series of units called vertebrae. the latter having the annelids and arthropods in being fundamentally metameric. however. on the other hand. which of these two divisions. As a rule. : All chordates have. but also a few primitive members that The presence may be referred to as Provcrtcbrata. and it was between distinction. as follows 1. With rare exceptions. They resemble Internally. but no additional skeletal structures are formed. externally as well. there are no external evidences of metamerism in the adult two openings. the vertebral column replaces the notochord. except in the lower chordates. once thought that all animals belong to either one or the other A few forms arc known. Thus the highest group in the animal kingdom -the Chord ata includes not only the Vcrtcbrata. arc a smaller group than either the arthropods or the mollusks but show a Like all the higher great advance over them in many ways. but in the lower fishes it 163 . numbering about 40. the chordates are bilaterally symmetrical. an unbroken rod In the provertebrates of supporting tissue called the notochord. possessing no true backbone.000 species.

The elongated body tapers at both ends and is It has a long median fin running along laterally compressed. These are clefts in the lateral walls of the pharynx. 115. never ventral to it as in the annelids and arthropods. being in reality not a "cord" but a tube. in its organization. and ending back of the anus. form is especially interesting in that it represents. around the tail.164 FUNDAMENTALS OF BIOLOGY No persists throughout life supplemented by the vertebrae. \ I ^ . and is invariably dorsal to the alimentary In canal. the vertebrates. permitting an outward passage of water from the mouth In the lung-breathing ehordates the gill cavity over the gills. In the gill-breathing forms these structures persist throughout life. They all first The agree. 115). From that point a pair of lateral dermal folds extends forward. in possessing basic chordate characters. 3. a possible ancestral stage through which the vertebrates may have passed in their early evolution. . It may swim rapidly through the water but usually lies buried in sand or mud near the shore. 2. The Provertebrates. being solely embryonic or larval structures. The three types of animals described below are very dissimilar in their general appearance and at first glance would seem to have nothing in common. fish-like animal about 2 inches long (Fig. All ehordates have paired gill slits at an early stage in their development. however. In ehordates the nerve cord is hollow. slits disappear early in life. the nerve cord is enclosed by the vertebral column. Dorsal fin Moufh Gill slit Myofome \ Fin rays Caudal Nerve cord Hofochord fin Pharynx FIG. The lancelet (Amphioxus) is a marine. the dorsal side. The Lancelet. X 2. animals other than ehordates have a notochord. Anus Derma! fold Sexual organs Airialpore\ Venfrat fin Afrium Hepatic caecum Infesfine Tho lancclot (Amphioxus lanceolatus) a primitive ohordate.

gs gill slits. No jaws are present. which is pierced by many pairs of gill slits. lancelet does not have a definite head. An impor- tant feature is the occurrence of paired nephridia. proboscis. after Ritter. Water enters the mouth and passes through the gill slits. p.THE VERTEBRATES The muscles The end in the 165 body are arrangement. t B up oxygen to the blood in the gills. lies just beneath the nerve cord and above the alimentary canal. . a chamber that partially encloses the pharynx. a sea squirt or tunicate (Cynthia). Agassiz. slightly enlarged. They wall are V-shaped and have a segmental known as myotomes. . B. It then enters the atrium. A. another sea squirt (Styela). C'. natural size. There is no skull and no brain. natural size. giving Primitive chordates. 116. c. extending the entire length of the body. (A. The mouth is located on the ventral surface a short distance from the anterior of the body. which terminates at the anus. The mouth opens into a sac-like pharynx. collar. the acorn-tongue' worm (Balanoylossua) FIG. and leaves through the atrial pore. The notochord. which open into the coeJom. after and C.) A. suggesting a remote affinity with the annelids. Food particles taken in through the mouth pass directly from the pharynx to the straight intestine.

Some are solitary. The head. pipiens). Four longitudinal nerve cords are present in the trunk. The poles. collar. arc degenerate be either free swimming or permanently Fi. outside. 116^4). the (Balanoglossus) is also found along seacoasts . nostrils. 117). Most of the features of a typical vertebrate may be seen in the frog. but no notochord is present. larvae are free-swimming creatures resembling small tadA short functional notochord is present in the tail. States. It is a worm-like animal having a proboscis. and trunk (Fig. The Acorn-tongue Worm. others colonial. The body is mostly sac-like and enclosed in a tough coat or tunic (Fig. -The bull frog (Rana catcsbiana). with gill slits. 117. or tunicates. Frog. larval stage that a relationship to other chordates is revealed. and a nervous system consisting of a It is chiefly in the single ganglion. attached to rocks and piles along seacoasts. the dorsal one being tubular and slightly larger than the others. chordates that The may sea squirts. Frogs are common inhabitants of marshes and other wet places. 116B and C). Water enters through a long upper tube and is expelled through a lower one. dorsal is to which a tubular nerve cord. a number of species occurring in the United The largest of these is the bull frog (Rana catesbiana.166 FUNDAMENTALS OF BIOLOGY worm Another primitive chordate. eyes. Fig. The Sea Squirts. trunk. one-half natural size. and eardrums are easily recognizable. easily recognized by. The larva of Balanoglossus is ciliated and motile and closely resembles the larvae of the echinoderms. There is a pharynx. but the leopard frog (R. acorn-tongue where it lies buried in the mud. The . It opens into a pharynx from which a number of paired gill slits communicate with the The notochord is small and located within the proboscis. The mouth is located between the proboscis and the collar. limbs. mouth.

the cloaca.walled. are facilitated by the presence of slime on the tongue and in the jumping and Digestive System. finally leaving the body through the anus. and tail. The head and a pair of eardrums. obviously an adaptation The short fore legs merely support the body. but the fore limbs four. bears a pair of large eyes. is the commonest species in the eastern and central parts of the country. and small insects. The vertebrate eye has a characteristic structure very different from that of mollusks or arthropods. The nostrils open into the large mouth Two jaws are present: an upper one firmly fastened to the cranium. scaleloss skin containing numerous mucuscavity. The body is covered with a smooth. are borne in a single mouth cavity. and mammals. but only the lowor one is movable. vertebrates are pentadaetyl having five digits (fingers and toes). The hind feet are webbed. None of the digits bears claws. The eye of the frog is provided with two eyelids.THE VERTEBRATES its 167 color pattern. Teeth row on the upper jaw and also in two localized groups on the roof of the mouth. products from the kidneys and the reproductive organs. The cloaca also receives vertebrates have a cloaca except all of the mammals. The teeth are small and all alike. but none occurs on the lower jaw. this being swallowed whole. trunk. amphibians. These are the fishes. The fleshy tongue. The frog belongs to the second of the five great classes of vertebrates. The extensive mouth cavity leads into a short esophagus. Then follows the long coiled small intestine and finally 118). The latter opens into a cavity termed the short large intestine. The mouth of the frog is very large. 117). External Features. sac-like stomach (Fig. a pair of nostrils. notched posteriorly. being used both in swimming. They are used not for chewing but merely for holding the food. but the frog has a tail only in early life (Fig. moist. reptiles. The body of a typical vertebrate is divided into head. which in typical secreting glands. which joins the thick. the thumb being present in a very rudimentary swimming. is attached at its forward end and is suddenly thrust out in capturing worms. in but the hind legs are long and powerful. where waste products of digestion accumulate. birds. All some of the fishes and almost . The capture and swallowing of food snails. Two pairs of limbs are present. for of the frog have five digits. The hind limbs have only condition. and a lower movable one.

The bile is carried from the gall bladder The to the anterior the bile duct. which form minute air sacs that open into a large undivided . 119). the three lobes of the liver The stomach has been moved to the Digestive system of the frog. wall (Fig. liver is the small green gall bladder. the secretes a digestive fluid into the small intestine through third gland should be mentioned. bile duct. red. and the small intestine spread out It pancreas. 118. turned back.168 FUNDAMENTALS OF BIOLOGY digestive system of the frog includes not only the digestive organs just mentioned. but also several digestive glands. a substance produced by the liver. which stores bile. globular body located near the point where the small and large intestine join. end of the small intestine by a slender tube called Another digestive gland present in the frog is the Gall bl. It is riot connected with the digestive system and has nothing to do with the work of digestion. A Respiratory System. The Attached to the largest of these is the red three-lobed liver. right. The frog has a pair of sac-like lungs lying in the anterior part of the coelom close to the dorsal body Their inner surface is increased by simple folds. the spleen a small. a white irregular body lying next to the stomach. to the left. Small intestine Liver Cloaca FIG.

frog. Meanwhile rhythmic contractions the throat force air of through the nostrils into mouth mucous membrane provides . Instead of being immediately expelled. of them chiefly by movements of the chest The circulatory system of a vertebrate Circulatory System. being kept there by closure of the glottis. veins. They are connected with the veins. addi- contional respiratory 1 J surface. The heart is enclosed in a sac called the pericardium. The lungs lie in the thoracic cavity. which pumps the blood. and. which is abundantly supplied with mucous glands and blood vessels. In the higher vertebrates a trachea (windpipe) extends from the glottis to the bronchial tubes. Capillaries and oxygen to the living tissues and remove waste products from them. The nostrils are then closed. T^ moist skin. ^ s 119. called the glottis.THE VERTEBRATES central cavity. through which it circulates. In man and other mammals the coelom is divided by a trans- verse muscular partition. These have very thin walls through which diffusion takes place. into a thoracic and an abdominal cavity. the larger ones giving rise to smaller and smaller branches that finally end in capillaries. This feature makes it possible for the frog to remain under water for a considerable length of time without having to come to the surface. by a contraction of the throat muscles. Thus vertebrates transfer nourishment . to a pair of bronchial tubes. Air is inhaled through the nostrils into the mouth cavity. the air is forced through a slit in the floor of the mouth. and capillaries. consists of a heart. and of arteries. which transport the blood back to the heart. there is In the lungs blood elaborate of an network vessels) capillaries (small through the walls of which oxygen is taken up and carbon dioxide given off. called the diaphragm. Ka ns of the the lun Respiratory orcomprising and bronchial tubes. siderable amount of respiratory gas exchange also takes place through the A _ FIG. one of which goes to each lung. Air is forced into and out and diaphragm. the air taken into the lungs of the frog remains for a considerable time. and out of the large where the moist cavity. Blood is carried away from the heart by the arteries. 169 The animal breathes with its mouth closed.

These are separated by valves so that the blood may flow only in one direction. Diagram of the circulatory system of the dogfish. forced forward through the ventral aorta. which gives off branches to the gills. which has thick muscular walls. 120). Only the larger blood auricle.} it is After passing into the ventricle. in contrast to the open system seen in the higher invertebrates. side view. Circulation in Amphibians. (After vessels are shown. Vertebrates that breathe with lungs have a more complicated circulatory system than the more primitive gill-breathing forms have. dorsal aorta. V. The heart of a fish consists of two fundamental chambers: an auricle and a ventricle (Fig. the veins. . For this reason we shall first consider the circulation of blood in a fish. where the blood flows out of the ends of vessels into sinuses. There it passes into capillaries where it absorbs oxygen from the water and liberates carbon dioxide. there being a left auricle. A. va. ventricle. Impure blood from all regions of the body is carried to the heart by the veins and is poured into the auricle. Alimentary fracf FIG. da. After it gives up its oxygen and food then returns to the heart through and collects waste products.170 FUNDAMENTALS OF BIOLOGY have a closed circulatory system. ventral aorta. a right auricle. From the gills the purified blood passes into arteries leading to the dorsal aorta. and consequently the blood makes a single circuit in going from the heart to the various organs and back again. The arrows indicate the direction of blood flow. giving off branches to all the principal organs. where it impure blood. 120. Wieman. The frog's heart is three-cham- bered. The most important feature to be kept in mind regarding circulation in the fishes is that the heart contains only passing into capillaries. Circulation in Fishes. which extends backward the entire length of the body. 121). and a ventricle The ventricle has much thicker walls than the (Fig.

amphibians have a This means that the blood makes two comin the circuits body. LA. chambered. 122). same direc- tion through the heart. It goes to heart is largely impure. then back to the heart. the veins carry blood to the heart. but here these comprise two sets: (1) systemic veins. aerated. evident that. which bring impure (non-aerated) blood to the heart from all parts of the body except the (aerated) blood fore limbs. Diagram of heart ventral view. and then the heart. body wall goes directly to the heart. from the systemic venous system. blood coming from the lungs through the pulmonary veins enters the auricle. cate the direction of the blood flow. goes to other parts of the trunk. the heart is The impure blood. and lungs. goes to It is double circulation. 121. it plete to the lungs. R A. left auricle. left From lungs Both auricles force blood time. through the pulmonary goes into the right ventricle walled and the ventricles thick walled. in contrast to the fishes. After leaving the various organs. finally to the various organs again.THE VERTEBRATES auricles. artery. and (2) pulmonary veins. consisting of two The auricles are thin two ventricles auricles and (Fig. which carry pure from the lungs. The arrows indiV. frog. into the ventricle at the same which then contracts before much mixing of pure and impure blood is Because the arterial syspossible. which is mixed. ventricle. skin to and be the lungs arises tem The next blood. Impure blood from the head. Returning to the heart . In the highest group of vertebrates. but that coming from other regions goes first either to the liver or to the kidneys. is largely pure and the head. four the mammals. to the lungs to be aerated. 171 As in the fishes. from the right side of the the first blood to leave the ventricle. as well as its constant flow in the the last blood right auricle. This disto goes directly of tribution blood. \ To lungs and arterial From systemic venous system The systemic veins empty their system Pure blood into the right auricle. is brought about by the presence of valves. poured into the right auricle and then. Circulation in Mammals. while of FIG.

In the frog.172 FUNDAMENTALS OF BIOLOGY through the pulmonary veins. pulmonary veins. and thence out through the aorta into the arterial system and to all parts of the body. left auricle. some of the veins. The bladder is a sac that stores the liquid waste products excreted earthworm. a. RA. pv. left ventricle. Kidneys are present in all vertebrates. 122. 126). ventral view. right ventricle. being a dilation of the ureters.A pair of oval dark-red kidneys lie in the coelom close to the dorsal body wall (Fig. RV. pa. the pure blood enters the left auricle. and one representing a higher state of evolution. Diagram showing the general scheme of circulation in a mammal. no mixing of pure and impure blood is possible. frog the bladder is a lateral outgrowth of the cloaca. Valves prevent the blood from Anterior flowing backward from the ventricles into the auricles and from the arteries into the ventricles. cause in the mammals left and right halves of the heart completely separated from each other. aorta. by the kidneys until ex- In the pelled through the anus. LV. The kidneys are connected with the cloaca by means of a pair of tubes called ureters. Thus it is evident that the mammals have are a more perfect scheme of circulation than the amphibians. passes into the left ventricle. but in the fishes and the higher vertebrates it is a very different organ. pass through the kidneys and give up nitrogenous waste products to them. pulmonary artery.. The arrows indicate the direction of blood flow. of tubules Each someof consists of a mass what resembling the nephridia the FIG. . right auricle. Excretory System. Be- the pulmocirculations are nary and systemic the because and entirely distinct. LA. in going to the heart.

" Ginn and Company. with right limbs removed. and Van Cleave.THE VERTEBRATES Skeletal System. size. the skeleton is entirely cartilaginous. 173 is The body internal framework of the frog (cndoskeleton) of bones supported by an cartilages (Fig. as in all vertebrates except the more primitive where Pectoral girdle Skull Hand Vertebral column Upper arm Uros+yle Pelvic girdle Foot Skeleton of frog. and spinal .) FIG. 123. The skeleton not only supports the body but provides places of attachment for muscles and protects such delicate organs as the eyes. Kelly. after Duges. fishes. and 123). brain. by permission. "General Zoology. natural (Redrawn from Linville. dorsal view.

forearm. the latter the two pairs of limbs and their" supporting limb girdles. such as occur in ribs. The skull includes the cranium. A. the jaws. ankle. or breastbone. A sternum. which. Each hind limb is made up of a thighj shank. (After Jewell model. The former includes the skull and vertebral column. is not joined to the vertebral column. and an appendicular skeleton. is composed of * * FIG. cranial nerves. / to X. the last pair being attached to the hip girdle. dorsal aspect. urostyle. and The vertebral certain other associated supporting structures. wrist. Brain of frog. ventral aspect.} nine separate vertebrae and a long posterior bone called the The urostyle is generally interpreted as a vestigial tail. Each fore limb consists of an upper arm. nearly all other vertebrates. is fastened to the shoulder girdle. or brain case. The fore limbs are .174 FUNDAMENTALS OF BIOLOGY The skeletal system of the frog consists of an axial skeleton cord. and a foot with five digits. and hand with four* digits. are lacking in the frog. B. unlike the hip girdle. 124. Each vertebra except the first bears a pair of short transverse True processes. column. serving as a supporting axis for the body.

the chief ones being a pair of olfactory lobes a pair of cerebral hemispheres. a cerebellum. (After Ecker. all vertebrates have a complex brain enclosed Netsal sac Olfactory nerve Cerebrum - Oph'c nerve 1st spinal nerve Sympa the tic system Spinal cord IO th spinal' FIG. and a medulla y oblongata (Fig. Although these parts are present in all . The brain has certain highly specialized parts. cranial and spinal nerves. and sympathetic nervous system.) by the cranium. 125. girdle. a pair of optic lobes. As contrasted with the invertebrates and lower chordates. ventral view. 124). the 175 hind limbs to the Nervous System.THE VERTEBRATES attached to the pectoral (shoulder) pelvic (hip) girdle. Central nervous system of the frog.

124). All the principal nerves have Kidney many The branches. by means of a number of slender sperm ducts the sperms do not pass through the kidneys and ureters. and consequently . Vertebrates have also a spinal cord.176 vertebrates. while in the mammals these parts are very large (Fig. while the spinal cord gives rise The to 10 pairs of spinal nerves. sympathetic nervous system. ^Bladder vertebrates have also a FIG. 126) Through these the sperms pass to enter the kidneys. In the higher vertebrates the excretory and reproductive ducts are separate. 125). male Excretory and reproductive systems of the frog. prises while the nerves that emerge Ureter from them. In the male frog a pair of small. yellow. FUNDAMENTALS OF BIOLOGY each group exhibits differences in its degree of development. 125). with all their branches. controlling their activities. oval testes lie beneath the kidneys and are connected with them (Fig. 126. Adrenal body central nervous system comthe brain and spinal cord. constitute the peripheral nervous system. chiefly the organs of digestion and circulation. Thus in the frog the cerebral hemispheres and cerebellum are relatively small. Reproductive System. The brain of the frog gives rise to 10 pairs of cranial nerves. latter innervate the skin Testis and the muscles of the trunk and limbs. which go to the sense organs and other important parts of the body. consisting of a chain of ganglia on spinal nerves (Fig. leaving the body by way of the ureters and the cloaca. In addiCloaca tion. is The location of the nerve cord above the in striking contrast to its position in the segmented worms and arthropods. a thick tube of nerve tissue extending backward from the brain and enclosed by the vertebral column enteron (Fig. each side of the spinal cord and connected with branches of the From these ganglia. nerves extend to the various internal organs. .

frog has a pair of irregular sac-like ovaries. . The lower end of each oviduct is dilated to form a distensible uterus where the eggs are temporarily stored. but because the mouth of the oviduct lies very close to the ovary. are not connected with the ovaries. through which the eggs leave the body. but open into the coelom. The eggs are liberated by Esophagus Open end of oviduct 'viduct Kidney Bladder ^xFio. Passing into the coelom. they find their way to the open ends of the oviducts.THE VERTEBRATES The female lie 177 below the kidneys but nearly eggs (Fig. As the eggs leave the body through the 1 In most of the higher vertebrates. 127). '. 1 oviducts they receive a gelatinous coating. chart. which fill the coelom when they are full of ripe The pair of long coiled oviducts. the oviducts similarly have no direct connection with the ovary. 127. which are funnelAs the eggs travel down the long coiled like and ciliated. the eggs do not remain in the coelom. ^ of the frog.) Cloaca Female reproductive system (After Pfurtschellcr wall a rupture of the ovarian wall.

128. Development. the embryo becomes a tadpole (Fig. 128). The eggs of the frog are always laid in water. gelatinous masses. mouth. When first hatched. and gills have not yet fully developed. which are enclosed by a fold of skin called the operculum. each mass containing thousands In about a week or 10 days after fertilization. for this reason it is thought that they contribute nourishment to the reproductive organs.178 FUNDAMENTALS OF BIOLOGY doaca. later they disappear and are replaced by internal gills. Stages in the development of the frog. 126 and 127). The first gills are external. Their exact function has not been accurately determined. 11 (From Wolcott. In both sexes finger-like fat bodies are attached to the anterior end of the sexual organs (Figs. Thus the embryos develop entirely outside the body.e frog. They occur in of eggs. the eyes. They contain a large in size amount of fat and always decrease during the breeding Oral sucker FIG. "Animal Biology. but these soon become visible. Water passes out of the . } season. they are fertilized by the mal.

and lateral paired fins are present. being about 3 months in the leopard frog and about 2 years in the bull frog. correspond to the fore and hind limbs.THE VERTEBRATES respiratory side. " A lateral " In cold-blooded animals the body temperature varies with the suranimals it is constant and relatively high. and tapered toward each end (Fig. is segments (myotomes). the To complete the metatail is slowly absorbed by the body. is external. Limbs are not developed until the is the organ of locomotion. but has a typical fish-like A long tail circulatory system with a two-chambered heart. in most cases. the intestine shortens. "warm-blooded" in roundings. THE GROUPS OF VERTEBRATES As contrasted with the provertebrates. The notochord. laterally compressed. In nearly all cases the skin is covered with scales. the of zigzag muscles in the body wall occur in the the seat of special sense organs. Fertilization. comprising the pectoral and pelvic fins. in man being normally 37 C. the mouth cavity enlarges. morphosis. the gills are replaced by lungs. numbering about Fishes. their appearance. movement through the water being accomplished by means of fins and a long flexible In the great majority of fishes both median unpaired fins tail. highly specialized for aquatic 15. and. The fishes. as both grow. are Typically the body is elong- ated. the heart becomes three chambered. which are metamerically line.). The duration of the tadpole stage varies with different species. arranged. then the fore limbs. There are no limbs present. The heart is two respectively. but in the lungfishes is incompletely three chambered. life. gills throughout life. persists throughout In all life in the lower fishes but is supplemented by vertebrae. all vertebrates have a cranium (brain case) and a complex brain.000 species. As form 1 in the lancelet. The latter. of the higher vertebrates. all of which finally result in the transformation of the tadpole into a frog. is or notochord other vertebrates the entirely replaced largely by a well developed vertebral column. and other changes take place.6 F. Fishes are cold-blooded 1 vertebrates having functional 129). chambered. 179 chamber through the spiracle. an opening on the left tadpole not only has gills. The Gradually the hind limbs make tadpole has reached its full size. (98. . which invariably appears in the embryo.

. The sharks and rays constitute an old group exhibiting many primitive features (Fig. only jaws but also paired fins. a large sac situated in the dorsal part of the coelom and concerned primarily with the maintenance of buoyancy. as follows: 1. a typical natural size. Another characteristic feature. A primitive feature is tho scales. The skin is smooth and without No air bladder is present. although noc a universal one. The tail is usually unequally lobed.180 FUNDAMENTALS OF BIOLOGs present in fishes. are not covered by an operculum.) bony fish. 129. but the notochord is slits The gill persistent and the skeleton entirely cartilaginous. 2. is the occurrence of an air bladder. cartilaginous. Elasmobranchs. they lack not to other fishes. The vertebral column and other parts of the skeleton are much more highly developed than in the cyclostomes. They differ so radically from the higher groups that they are generally regarded not as true fishes but as forming a class by themselves. one-half funnel-shaped mouth by means of which they attach themselves In contrast to the higher groups. above which but not enclosing The skeleton is entirely it lie the poorly developed vertebrae. Modern fishes are of five Cyclostomes. DormI Caudal fin FIG. The cyclostomes are parasites fins with a jawless. The yellow perch (Perca Jlavescens]. as they are in all the higher fishes. The mouth is ventral and situated a short distance from the anterior end of the body. -The most primitive fishes constitute a small group comprising the lampreys and hagfishes (Fig. ISO/I). 130#). presence of a persistent notochord. (After Forbes. chief types.

spiny dogfish (Squalus acanihias). Fishes of diverse types. 130. sea lamprey (Petromyzon marinus). primitive type. J5. a ganoid. no air bladder is present. the vertebral column extending into the upper The scales are of a lobe. A. after Dean. D. after Gunther. after Jordan. . common sturgeon (Acipenser sturio). each consisting of a bony plate bearing a toothIn contrast to the higher fishes. FIG. The sturgeons resemble the elasmobranchs in having a ventral mouth and a heterocercal tail. are repre3. also an old group. a shark. sented chiefly by the sturgeons and gar pikes (Fig.THE VERTEBRATES 181 or heterocercal. D.) The ganoid fishes. A. Australian lungfish (Neoceratodus miolepis). which is larger than the lower one. 130C). C. a cyclostome. Ganoids. but in the gar pikes the mouth is terminal and the tail either heterocercal or homocercal (see next page). The scales are typically rhomboidal and enameled. but that of the gar pikes is more completely so. B and C. like spine. The skeleton of the sturgeons is only partly ossified.

Nearly all amphibians have entirely which are four limbs typically pentadactyl. but the skeleton is partly ossified. Fertilization.182 FUNDAMENTALS OF BIOLOGY These are the bony fishes. This class includes only about 2. The heart is and one ventricle. Historically. The scales are 5. the muscles are arranged in longitudinal bands rather than in zigzag segments. without scales. with rare exceptions. but the air bladder opens into the pharynx and functions as a true lung. 130Z)). 129). out their entire existence. cereal. round and overlapping. They are thought to have arisen from some ancient fish stock. diversified in the past They were more abundant and than they are now.000 species. generalized 1. The scales are round and overlapping. but not directly from the lungfishes. The skeleton is composed largely of bone. group including over 95 per cent of all existing species (Fig. As a rule. the heart thus being incompletely three chambered. forms with an elongated body and a persistent tail (Fig. and the animal adopts a terrestrial mode of group is that. Salamanders and Newts. stagnant pools and marshes. As in the clasmobranchs the notochord is persistent. South Australia. The mouth is terminal. but in most cases the gills are later replaced by lungs. the great modern 4. The tail is the upper and usually outwardly symmetrical. and Brazil (Fig. There are two chief groups of modern amphibians: These are primitive. Africa. the toes being without life. once but now represented by only three genera confined to abundant. the gills are persistent and functional throughout life although small lungs . In such forms as Necturus. there hoing two auricles most cases. two pairs of limbs are present. All the lungfishes have gills. A unique feature of the is the skin claws. They live in Lungfishes. is external and the eggs are laid in the water. 131). As in all the higher groups. The lurigfishes are an ancient group. Teleosts. Some retain their gills arid remain aquatic through- Amphibians. or homolower lobes being equally prominent. the amphibians are an old group. but the posterior pair is sometimes wanting. in three chambered. An advanced feature of the group is the presence of a partial partition in the auricle. there are approxi- mately equal in size. are cold-blooded vertebrates with functional gills Amphibians in early life. the common mucf'puppy.

water. or both. Toads and Frogs. FIG. The heart is three chambered. Like the two lower but show a functional without at any in advance gills entirely being great of the in Some members live of group development. "Animal Biology") . stage but all reptiles have functional lungs and so breathe air. as in most of the classes. 117). mud puppy (N ccturus maculosus) X H?^ larva and C adult of tiger salamander (Ambystoma liyriniim). (From Wolcott.000 species of living reptiles. Reptiles. In contrast to the amphibians. A. higher vertebrates. Salamanders. and the toes end in claws. in adult life 2. the skin is covered with scales. X %. 183 gills In many other forms the are replaced functional lungs. a notable exception).THE VERTEBRATES are also developed. Two pairs of limbs are always present. the hind limbs being much Gills are present only in early life. In these amphibians the by body is short and the adult stage of development (Fig. larger than the fore limbs. tail in broad and lacks a in all cases being later replaced by functional lungs. y . There arc about 4. hard Nearly all reptiles have four limbs (snakes being plates. reptiles are cold blooded. as in the . 131.

3. even those of aquatic reptiles. A. Teeth are present. Teeth are absent. There are only three main surviving groups: FIG.184 FUNDAMENTALS OF BIOLOGY amphibians. 132. Like the amphibians. a lizard (Gerrhonotus scincicauda) tortoise (Testudo agassizi. In all snakes arid a few Teeth are present. limbs are entirely lacking. but the ventricle is partially divided. In the crocodiles and alligators. Scales may be present on other parts of the body. lizards. The heart is almost completely four chambered. The shell is composed of bony ened to the vertebrae and ribs. . The body of these forms is elongated and covered with hard scales that do not overlap. The eggs are always laid on land. Crocodiles and Alligators. which hatch outside the body. reptiles have had a long geologic history and were a much more important group today. X J. X > desort These reptiles have an elongated body 1. Reptiles. . These are reptiles with a short body plates firmly fast- enclosed within a shell consisting of an upper and a lower portion (Fig. 132^4). but in most cases shelled eggs are laid. covered only with scales (Fig. Lizards and Snakes. in the past It is believed that reptiles arose ages ago and later gave rise to both birds than they are from amphibians long and mammals. Fertilization is internal without exception. however. sometimes it is covered also with bony plates. Turtles and Tortoises. 1325). the heart is almost completely four chambered. 2.

The skin is covered with feathers. and swans are also toes (Fig. enclosed by a horny bill. mentioned here. which hatch the presence outside the body. conserve the heat of the body and increase the surface of the wings In all birds the fore limbs are modified to flight. heart is The completely four chambered. 133. They live mostly on fishes. 133). as in reptiles. All birds lay shelled eggs. The legs are placed far back on the body. Feathers and tail in form wings. The gulls and terns are long2. The jaws. they lack gills at all stages of development. auks. 2075).THE VERTEBRATES Birds. Long-winged Swimmers. geese. The orders of modern birds are so numerous that only some of them can be briefly 1. Anserine Birds.000 species of birds form a very distinct class Birds are specialized for aerial life (Fig. so that a more maintained on land. swimming webbed The beak is . The loons. of the Fertilization is internal. Diving Birds. or less erect attitude is 3. Like the reptiles. but show an advance in being warm blooded. scales occurring only on the feet. birds with The ducks. winged water birds with sharp-pointed or hooked beaks and webbed toes. and penguins are diving birds with short wings and webbed or lobed toes (Fig. 185 The of vertebrates. Birds are the internal organs and often inside the bones closely related to the reptiles and were derived from them during middle geologic time. Crow (Corvus brachyrhynchoa. A special feature group is FIG. 207C). have no teeth. 14. grebes. X of air spaces among as well.

. These are scratching birds. hooked beak for tearing flesh.0. 8. 206(7). all They are also known our common songsters. 133. are nearly always present. four webless toes of equal length arranged on the same and one behind as (Figs. pointed. Shore avocets. sparrows. blackbirds. 7. the hind one fowls (Fig. The woodpeckers and flickers are climbing birds having sharp-clawed toes usually arranged with two in The bill is straight. of all known and have T hey are of small or medium size species of birds. bills Raptorial Birds. quail. slender. stilts. turkeys. plovers. 134). 4. stout. vultures. being lifted above the ground. strong. convex 6. 207/1). and very insects. Birds. eagles. etc. " " warblers. Two pairs of limbs Mammals. mammals are warm blooded and at no time have gills. arid more getting food from mud.00 species. and domestic The toes are short and stout. crows. They have long slender legs adapted for wading and an elongated beak used in The toes are long. 206). The powerful clawed feet are adapted for seizing living prey. group is the fact that the young are nourished by milk secreted bv the mammary glands of the female. or less lobed. are small or medium-sized birds common along the margins of streams and lakes (Fig. the stout. sandpipers. thrushes. 206/1). stout. young undergo their early A distinctive feature of the development within the mother's body. This order includes approximately half level with three in front and 206D and E). on carrion. Gallinaceous Birds. This order comprises the birds of prey. 5. and many others. ^The vultures live Woodpeckers. which may be reduced in area or nearly wanting (Fig. pheasants. The skin is covered with hair. perching birds and include examples being the flycatchers. front and two behind (Fig. and owls (Fig. such as hawks. including grouse. The heart is completely four chamFertilization is internal. These birds have short. and in almost all mammals the bered. swallows.. partridges. being used for digging into bark and wood for Passerine Birds.186 flattened FUNDAMENTALS OF BIOLOGY and often furnished with tooth-like projections that act as strainers for food. The 5. adapted for crushing seeds. highest group of vertebrates numbers about Like the birds. finches. wrens. The snipes.

the marsupium. The more important mammalian groups are presented below: 1. 208 and 215). They are also characterized fact that the mammary many The Marsupials. inside of These arc the most primitive of the placental mammals (see p. Many of the inscctivores are burrowing in their habits. 134. X and 2. A large web extends between the greatly lengthened digits of . and are differentiated into four distinct typos. simple. 240) and include the shrews. The monotremes are the most primitive of The duckbill and spiny ant eater of Australia and adjacent islands are the only surviving representatives (Fig. and the opossums of America. and hedgehogs (Figs. its birds. They are small creatures that eat Their teeth are small and insects and other small animals. The Australian relatives. existing mammals. young are born in a very immature state and are placed by a cloaca and by the glands are not localized. moles. Monotrcmcs. Mammals originated from reptiles in middle geologic time or somewhat earlier but did not reach their greatest development until within relatively recent times. 4. Ground squirrel (Citellus beecheyi). Insectivores.THE VERTEBRATES 187 The development of the brain far surpasses that seen in any of In most cases the teeth develop in two sets the lower groups. These curious forms lay shelled eggs like those of reptiles 163). Bats. marsupials comprise the kangaroo and within an abdominal pouch called which are the mammary glands. FIG. The bats are the only mammals that really fly. 3.

Sirenians. Bears. prairie dogs. The rodents are small mammals but exceed all 6. food.188 FUNDAMENTALS OF BIOLOGY the fore limb. skunks. The teeth of carnivores are adapted for tearing and cutting flesh (Fig. The even-toed ungulates elk. and the dugongs of Ocean. comprising the . while the fore limbs are paddle-like. Ungulates. (artiodactyls) include the cattle. to a diet consisting of fruit or of insects. sheep. These are the hoofed mammals. which walk 8. arid hogs. . Elephants. whales. They are sluggish aquatic upon plants. South America bite into the skin exuding blood. one native of Africa. antelopes. squirrels. the Indian of the Atlantic Australia. porcupines. dogs. 213C). As in the last group. hairy anteaters. armadillos. 5. They include the rats. There are only two living species of elephants. otters. 218C) the hind limbs are wanting. This group includes the manatees or sea cows 10. 134 and 210). camels. Rodents. ? These are the flesh-eating mammals. cats. Coast of America and Africa. 212). weasels. Edentates. forming a wing (Fig. 217). The of the victim vampire bats of and lick up the Carnivores." The whales are the largest living mammals. This is also an aquatic order. The horses. raccoons. and scaly anteatcrs. gophers. their teeth 'being adapted for gnawing (Figs. teeth of ungulates are adapted to plant food (Fig. beavers. mice. reaching to the sides of the body and to the hind The simple teeth are adapted limbs. on the tips of their toes (Fig. while the seals sea lions. hippopotamuses. the fore limbs and tail being paddlelike.$ hoofed. hares. and the Red Sea. guinea pigs. The nose is developed a prehensile trunk. 211). 9. is functional toes and each toe a. The feet have five tapirs. deer. The and rhinoceroses are odd-toed ungulates (perissodactyls). and rounded mammals living along seacoasts and feeding The hind limbs are absent. This order includes mammals characterized by an absence or imperfect development of teeth. dolphins. Cetaceans. tail 11. and badgers are and walruses are marine representatives (Fig. 7. and the two upper front teeth as tusks. the other groups in number of species. the other of Tndia. terrestrial forms. arid porpoises (Fig. Some cetaceans have but in others teeth are replaced by "whalebone. teeth. 216). Hair is almost completely absent. such as the sloths. goats. etc. giraffes. For the most part rodents eat plant rabbits.

209 and 213^4). Primates. 189 The primates. Physically they only Primates are simpler than most of the other mammalian groups. comprising the lemurs. the of on basis their mental development. The digits have nails rather than claws or hoofs. are prevailingly arboreal forms with relatively primitive teeth and limbs (Figs. . are and man entitled to first place in the animal kingdom apes.THE VERTEBRATES 12. monkeys.

stance called mucus. It includes the outer part of the skin (epidermis) and the lining of the various internal cavities and tubes.CHAPTER XI CHIEF ANIMAL TISSUES A tissue is a group of similarly differentiated cells performing one or more functions in common. absorption. The sponges and or of cells and products derived from them. all of which arise from undifferentiated cells of the embryo. The mucous consist of epithelial tissue. as in the outer portion of the comprising human skin. many layers. Its cells are relatively small. Epithelial cells what modified may cells. but in members of the higher metazoan groups complex tissues of a number of different kinds are present. various types of epithelia are present. which respiratory and alimentary tracts. which may be somein accordance with the functions they perform. coelenterates are characterized by relatively simple tissues. nerve tissue^ and connective and supporting tissues. In the sponges and coelenterates the body is composed almost entirely of epithelial cells. and compactly arranged. muscle tissue. on the other slightly specialized. 135). regular. Each of these will be briefly considered. Epithelium. it is stratified. reproductive cells. and are often ciliated Epithelium commonly consists of a single layer of but often. flat. Mucous membranes. most of which are but In more complex metazoans. hand. line the They secrete a sticky sub- membrane of the respiratory passages is composed of a single layer of columnar epithelial cells with cilia. all the tissues of the With the exception of the body of the higher animals may be referred to four main types: epithelium. This is always a lelatively simple type of tissue that covers the external and internal surfaces of organs and forms the secreting cells of glands. or columnar. cubical. be either (Fig. The beating of the cilia tends to sweep bacteria and other foreign bodies out of the air passages. secretion. such as protection. It may consist only of cells. The mucous mem190 . etc.

") abdominal cavity and covers the abdominal organs. X 300. and small intestine. D. (From Wieman. the pleurae line the thoracic cavity and cover the lungs. For example. ciliated epithelium from pharynx. surface view of flat epithelium from skin. 135. and the glands of the stomach. C. B. cubical epithelium from kidney tubule. "General Zoology. abundant in invertebrates and represented in man by the scattered goblet cells that occur in the intestines (Fig. A. epithelium. They also cover the organs lying within and serve as a means of attachment for them. Examples are the sweat glands and oil glands of the skin. mucous membrane lining The various other . The simplest glands are one-celled structures. the cavities that do not communicate with the surface of the body. the stomach and 136). faciliSerous membranes line tating the onward movement of food. The mucus secreted by it serves as a lubricant. Several types of epithelial tissue from larval salamander. pancreas. Glands are composed chiefly of modified epithelium and secrete substances of various kinds. while the pericardium forms a covering for the heart. columnar epithelium from intestine.CHIEF ANIMAL TISSUES brane that lines 191 the alimentary canal consists of stratified . the peritoneum lines the FIG.

uninucleate cells pointed at either end. such as those of the walls of . C. 136. simple alveolar gland. hoofs. In oil glands and milk glands. B. The secreting portions of the glands are stippled. They compose the muscles of most of the lower animals and nearly all the involuntary muscles of the higher animals. or tubular and may be either branched or unbranched. flask-shaped. compound saccular gland. Muscle Tissue. They may be cup-like. the secreting E F t FIG. A branched gland is The secretion said to be compound. claws. 137). usually leaving the gland cells intact. is poured into the cavity of the gland. they are replaced substance called keratin. broken down and pass out of the gland with their prodby newly formed cells. Z). simple saccular gland. A. Nails. and horns are composed of compact layers of epithelial cells that have become greatly hardened with a cells are uct. F. composed of cells that are specialThere are two kinds of muscle: smooth and striated. an unbranched one simple. Smooth muscle consists of layers of elongated. however. Diagram showing types of glands. E compound tubular gland. simple tubular gland. Muscle is ized for contraction (Fig. goblet cells.192 FUNDAMENTALS OF BIOLOGY glands are many celled and form depressions lined with epithelium.

Striated muscle FIG. 137. The fibers of each nerve cell are nearly always of two kinds: dendrites. greater and body from which extend thread-like cytoplasmic fibers that carry the nervous impulses (Fig. blood vessels. They are made up of bundles of long fibers with numerous transverse striations. and an axon. it is involuntary. is is enclosed by a layer of connective highly specialized and capable of much while each bundle of fibers tissue. smooth muscle cells. which normally cell carry impulses toward the cell body. Its functions are to receive and conduct sensations and to stimuA nerve cell consists of a nucleated late activity in other cells. with two nuclei. trachea. each fiber representing a row of cells that have fused together.CHIEF ANIMAL TISSUES 193 the digestive tract. uterus. Nerve Tissue. Voluntary muscles. A. and of certain glands and their ducts. 138). which carries them away. Generally the dendrites are relatively short and . which are those under the control of the will. Each fiber is Scattered nuclei occur at intervals along the surrounded by a firm membranous sheath. more rapid contraction than smooth muscle. part of a striated muscle fiber. are striated. fiber. This kind of tissue is also highly specialized. B. bronchial tubes. although striated. Muscle tissue. bladder. The muscle tissue that forms the wall of the heart is of a special type in that.

Connective Tissues. tip. cell body. are said to be medullated. The fibers may occur singly or in bundles and are unbranched. cartilage. A ganglion is an aggregation of cell bodies. axon. d. are surrounded Most nerves of by a sheath connective tissue. 138. tissues arc characterized by the ms presence of some kind of intercellular substance that is mostly non-living itself but is nearly always formed as a secretion from the p living cells. It consists of FIG. determines the character and functions of the tissue. c. rows of flattened cells distributed throughout a gelatinous matrix containing. rather than the living cells. Diagram of a typical a. They rigidity include fibrous tissue.delicate white fibers. etc. ma. while those which lack a sheath are non-medullated. as well as the cover- . while a nerve is a bundle of nerve Some nerve fibers have a sheath of fatty material and fibers. bone. White fibrous connective tissue is non-elastic. dendrite. but are specialized. The matrix. sequently the cells are Conemfluid. while the axon is very long and branched only near its A nerve cell with its fibers is called a neuron. White fibrous tissue is tough and flexible. riety of to bind and Supporting These include a vatissues whose function together various body or to give is parts of the and support. medullary sheath. It forms the tendons and ligaments. All these. blood. neuron. bedded in themselves matrix (or a little in the case of blood). It is found in many parts of the body.194 FUNDAMENTALS OF BIOLOGY branched.

especially in the walls of arteries lungs. Elastic tissue is found throughout the body. X 500. 140." Longmans. line cartilage. Yellow fibrous connective tissue is made up chiefly of many elastic fibers. of Cross section of bone. the fibers may be either white or yellow. while ligaments are bands that hold bones together. The in cells are numerous and crowded together. depending on the character of the matrix surrounding the cells. FIG. 139. Fatty (or adipose) tissue consists of cells containing large globules of fat and surrounded by a matrix of white fibers and a few elastic fibers. the bone cells lying in the smaller " Textbook of (From Schafer spaces arranged in concentric circles. X 150. but branch extensively and tend to form a net- work. Hyaline cartilage has a .) largfc Cartilage may be either fibrous or hyaline. by permission. the intercellular substance in white. Microscopic Anatomy. The FIG. after Sharpey. Green & Company. Fatty tissue is abundant bone marrow. It also occurs in layers in the deeper parts the skin and around various internal organs. ings of muscles. They usually occur singly.CHIEF ANIMAL TISSUES 195 Tendons serve to attach muscles to bones. and in the walls of the It also occurs as shock-absorb- ing pads between the vertebrae. If fibrous. Section of hyashowing the cells imbedded in a non-living matrix. black areas are occupied by blood vessels.

and 33 per cent The bone cells lie in small cavities connected organic matter. It covers the ends of bones. and fat cells. The cartilage cells the chief supporting tissue of vertebrate animals. bone consists of about 58 per cent calcium phosphate. It is composed of cells that secrete a hard matrix of mineral matter.196 FUNDAMENTALS OF BIOLOGY any fibers but actually with very occupy small spaces in the matrix. The the white corpuscles (Fig. The small cavities containing the bone cells are arranged in concentric circles that surround larger canals occupied by blood vessels. and bronchial tubes. connective tissue. and other connective tissues the intercellular substance duced by the cells. 141). X 750. which is generally laid Human in the form of thin concentric layers (Fig. FIG. larynx. joins Cartilage is firm and tough. clear matrix apparently without small ones. The bone is surrounded by a connective-tissue membrane that supplies it with nerves and blood vessels. 140). certain bones together. each space containing one or several cells (Fig. ear. The shafts of long bones contain a marrow cavity. 2 per cent other mineral salts. Blood consists of cells that float freely in a liquid. 139). is Bone down consisting chiefly of lime and phosphorus. 7 per cent calcium carbonate.^Blood corpuscles of a frog. trachea. with one another by delicate cytoplasmic strands that penetrate the matrix. . but unlike is not prothree principal constituents of the blood of vertebrates are the liquid plasma^ the red corpuscles. four of them red and two white. 141. and forms the supporting tissue for the nose.

. expending a vastly greater amount of energy. We present interest is not so much in the nature of these metabolic processes as in the ways in which they are carried on in animal bodies. VII). fats. and how it is formed in green plants. of complex digestive and excretory organs. and proteins and that these are utilized by both plants and animals in exactly the same ways. they have a much higher rate of metabolism. the greater part of the "food consumed by animals is used to replenish energy expended in muscular movements. For this reason most of the ordinary activities of animals are concerned primarily with the procuring arid utilization of food. tion. Because animals are active and perform their functions in a more accelerated manner than plants. Animals show great variability with respect tp the kinds of materials upon which they feed and to their 197 . necessitating in most cases the development of special prehensile organs. in the building up of body substance (both protoplasm and its derivatives) and as a source of have seen that the processes of digestion. of special sense organs. especially in those of the higher animals. We have seen that all organisms use the same kinds of foods carbohydrates. Feeding Habits. arid of many other complexities in organization and behavior that green plants do not have.CHAPTER METABOLISM IN XII ANIMALS Through our study of plants wo have become acquainted with a number of aspects of metabolism that are fundamental to all We have learned what food is living things (see Chap. energy. The distinctive feature of nutrition in animals is their inability to carry on photosynthesis and their consequent dependence for energy and formative material. of a means of locomotion. upon food previously made by green plants. viz. and demanding a correspondingly larger consumption of food. In fact. assimilaand their accompanying energy relations are Therefore our essentially similar in both plants and animals. either directly or indirectly. respiration.

Fats are obtained chiefly from fatty meat. Dietary Requirements. fats. when oxidized in the body. eggs. 15 per cent. 1 They are of greatest value in the building up of tissues and for this purpose are When digested. calories per gram. Foods rich in proteins include lean meat. proteins are broken down into indispensable. milk. cereals. as to tapeworms. Because fats contain relatively less oxygen than do carbohydrates. same amount porated into the body. fats. etc. eggs. in about the following proportions: carbohydrates. animals require for normal growth and proper functioning of the body adequate quantities of certain accessory food materials. or to only one type. 65 per cent. and vitamins. examples being vultures and hyenas. capturing living animals and devouring them. fats. while carnivorous forms Some carnivores. living either on the outside of the body. which are then rebuilt into new proteins and incor- more energy. whole-grain cereals. of food. In addition to carbohydrates. and proteins. are eat animal food. These Water is not only an are water. yield the of energy as carbohydrates. constitutes about 65 per cent of the weight of the human is The heat produced by fats. oxidized in the body. or inside. such as hawks and wolves. and salad oils. milk (especially cream and butter). these are their only true food substances. while carbohydrates and proteins yield only 4 . predatory. and proteins furnish the only sources of energy that animals utilize. 20 per cent. A well-balanced diet would include the three main classes of food substances. inorganic salts. Herbivorous animals feed upon plants.198 FUNDAMENTALS OF BIOLOGY that means of obtaining their food. based on weight. eating carrion. Parasites attach themselves and extract nourishment from other living animals. but the diet of most animals is restricted to a few types. as fleas and lice. Man is omnivorous. and beans. amino acids. proteins. milk. others are scavengers. they are capable of undergoing greater oxidation and so yield Proteins. the The chief sources of carbohydrates in human diet are potatoes. Water body. equal to 9 calories. dung. Some animals live on plant or animal products. properly speaking. peas. using foods come from many different sources. Since carbohydrates. such as nectar. and sugar. dead leaves. essential constituent of protoplasm but a necessary solvent and a medium 1 in which chemical reactions when may be carried on. bread.

beri- and pellagra are definitely known to result from a lack of specific vitamins and so are appropriately termed " Each of these can be entirely prevented deficiency diseases. C. winter squash. particularly of milk. have been isolated in pure chemical form. as constituents of certain digestive juices. carrots." or effectively cured proper vitamins. and white bread. a substance present in all green plants as a constituent of chloroIt also occurs in many orange or yellow vegetables and phyll. This vitamin is derived from carotin (C-ioHBe). dairy products. An insufficiency of vitamin A .METABOLISM IN ANIMALS Mineral 199 salts are essential to the development of bones and teeth. Vitamin A. egg yolk. scurvy. an adequate supply of each of the necessary vitamins will be obtained if the diet is sufficiently diversified to include wholevariety and quantity. At the present time six different kinds of vitamins are recognized. Foods cots. fruits. Vitamins A. vitamins B. and today much progress is being made toward a determination of the vitamins least four of the exact role that each plays in human metabolism. to the regulation of nervous activity and the control of muscular contractions. vegetables. particularly important to the maintenance of health and the proper growth of infants and children that their diet contain the necessary vitamins in sufficient amounts. beri. Some and at have been synthesized. and fruits. sweet potatoes. by eating foods containing enough of the Although some foods are deficient in vitamins. grain cereals. Vitamins are organic substances present in many Vitamins. An insufficient quantity of any one of these in a diet wise adequate causes that is otherIt is marked functional disturbances. rich in carotin or in vitamin liver. eggs. as to both especially meats. and G are soluble in water. foods of both vegetable and animal origin. in the prevention of an acid condition of the tissues. and E are soluble in fats. of our and fruits. A (CaoHaoO). rickets. The six different vitamins have been designated by letters. Most knowledge of vitamins has been obtained in recent years. very small amounts of which are necessary to the normal functioning of the body. and all A include butterfat. aprigreen vegetables. Salts are constituents of most foods. etc. vegetables. D. cod-liver oil. When is the carotin plant food containing carotin is eaten converted in the liver to vitamin by animals. Xerophthalmia. potatoes.

a substance produced when ergosterol is exposed to ultraviolet radiation. is especially abundant in yeast. and loose teeth. It is also present in unpasteurized milk. liver. to wheat germ. spinach. irradiated milk. Ergosterol is present in many fat-containing foods which. which is cevitamic acid (CeHsOe). body One of its forms is viosterol (C2sH43OH). such as white flour. Vitamin D. polished rice. and When the skin is exposed to ultraviolet light. . in the respiratory and digestive susceptibility growth.200 in the FUNDAMENTALS OF BIOLOGY body causes abnormal changes tracts. Vitamin C. and hominy. loss of appetite and weight. digestive disorders. and a disease of the nervous system known as beriberi result from a deficiency of vitamin B in the diet. parsnips. when irradiated. A deficient amount of this vitamin in the diet causes scurvy. irradiated yeast. is abundant in oranges and other citrus fruits. retardation of eye disease called xerophthalmia. This vitamin. peanuts. milk. bleeding. by defective bone structure. fruits infections. but this has not been This is established. grain hulls. Tooth decay and certain gum infections be due to vitamin-C deficiency. A substance identified as thiamine (Ci 2 Hi 8 N 4 OS). Rickets in infants and young children is a manifestation of faulty calcium and phosphorus metabolism. is either from the sun or from an artificial source. Vitamin to a considerable extent this is and cabbage. a term applied to one or more substances and phosphorus properly. and an Vitamin B. was once common among sailors whose diet was lacking in by heating in the presence of oxygen. such as tomatoes. It is deficient in artificially refined foods. Retarded growth. fruits may and vegetables. as well as in certain vegetables. tender gums. and It is also important in tooth corrects prevents in a sound tooth structure. eggs. vitamin vitamin include cod-liver D formed within the body. C is destroyed but avoided in commercial canning operations. egg yolk. and often designated as vitamin Bi. Beriberi occurs chiefly among Oriental people whose food consists largely of polished rice. Foods rich in this furnish an important source of vitamin D. that enable the to utilize calcium oil. a disease characIt terized by weakness. Vitamin D this disease. night blindness. and many and vegetables. although formation and maintaining there is no convincing evidence that an adequate ajnount of It is characterized vitamin D in the diet will prevent tooth decay.

and large intestine. a substance occurring in fairly high concentration in whole-grain cereals. stomach. which is often designated as vitamin 62. small intestine. digestion is intracellular. coelenterates. Its absence may be responsible for infertility. it is carried on inside individual cells. most of which are insoluble. of vitamin G results in retarded growth and impaired health. liver. a That of a number of differdefinite digestive system is present. digestion in animals is accomplished by enzymes. yeast. tive cavity. There is some confusion in regard to the identity of this vitamin. The coelenterates and flatworms also carry on extracellular digestion. disturbances. eggs. Here food enters and is digested within a distinct diges- unknown chemical composition Enzymes are secreted into the cavity. sponges. at least in some cases. Some of the foods taken into the animal body. must be broken down into simpler substances that can pass into solution. Digestion. although it has been supposed by some to be vitamin G. but its need in the human diet has not been established. Vitamin G. and nervous disorders. and a few ent animals except the protozoans. such as simple sugars. Com- plex foods. parasitic forms belonging to the higher groups. As in plants. can be absorbed directly. but most nutritive materials must undergo chemical change before they can be utilized. substances of that the organism itself produces. The digestive system of man and other vertebrates consists of a mouth.METABOLISM IN ANIMALS Vitamin E. the pellagra-preventing factor. or by accessory digestive glands connected with it by ducts. and green leafy vegetables. and to a certain extent in the coelenterates and flatworms. that is. yeast. A deficiency milk. either by the surrounding cells. The purpose of digestion is to render foods capable of being absorbed and ultimately assimilated by living tissues. It appears to be the same as lactoflavin (CnH^oN^). all In animals has been described in previous chapters. and this is the method characteristic of all the higher groups. . digestive It is of dietary origin and be vitamin a may prevented by present in wheat germ. each cell engulfing its own food. and There is considerable uncertainty as to the identity of liver. esophagus. 201 A vitamin present in wheat germ and in lettuce is essential to rats for successful reproduction. In the protozoans and sponges. Pellagra is a disease characterized by skin lesions.

202 FUNDAMENTALS OF BIOLOGY These organs form a continuous tube. FIG. ie can i nes or eyeteeth. there being in each jaw 4 incisors. the liver and pancreas. and the root the basal portion The crown lying in the jawbone. Diagrammatic lonsection of a human gum. It lies just beneath the diaphragm. set. intestine arises at the lower end of the stomach. a substance softer . in the adult. the alimentary canal. 142). tooth consists of a crown. teeth. is tooth made of dentine. A duct coming from the liver joins the pancreatic duct near its lower end to form a common duct that enters the small intestine at a point about 4 inches beyond its origin at the end of the stomach. and The crown is the portion projecting beyond the gum. the bile duct. the neck the part lying in the gitudinal tooth. extending The small transversely across the body from left to right. 4 pre- Enamel Dentine molarSj and 6 molars. which arc broad and premolars. both of which are very hard. which leads to the gall The lower bladder. It is a narrow. of The flat. and tongue are taking food into the mouth and in preparing it for swallowing. or throat cavity. to which are attached two main accessory digestive glands. Man has. molars. similar and the form but Gum Pulp cav'+y smaller. The mouth leads into the pharynx. while the chisel-shaped incisors are adapted for biting and cutting. highly coiled tube about 23 feet in length. are conical ^ A > and are used for biting and tear- Cement Uctwbone ing. are used for grinding food. neck. which is connected with the stomach by means of the long tubular esophagus (Fig. The greater part of the than enamel. The duct of the liver has a branch. The lips. The stomach is a muscular sac about 11 inches long and with a normal capacity of about one to one and a half quarts. 143). In the center is the pulp cavity containing nerves and blood vessels. 2 canines. root (Fig. is covered with enamel and the root with cement. 32 teeth. a sac in which bile is temporarily stored. Human used in Digestive System. 142.

but lie pairs are present. stomach. The largo intestine. from right to left just beneath the stomach. are the ascending colon. and an enzyme. R. pancreas. 143. sigmoid flexure. 0. S. It first passes upward. of greater diameter than the small intestine. then across the body is about 5 feet long. small intestine. which serves as a lubricant. transverse colon. leaving a blind sac. $1. after making an S-shaped curve. liver. descending colon. TC.METABOLISM IN ANIMALS end of the small intestine 203 opens into the large intestine a short distance from its end. Digestion in Man. in the order described. where the food saliva. AC. The different parts of the large BD- SF FIG. the caecunij to which is attached the slender vermiform appendix. vermiform appendix. of which three These glands are not in the mouth. caecum. continues as a straight short tube to the anus. DC. and rectum. duodenum. transverse colon. descending colon. Diagram of human digestive system. L. sigmoid (From flexure. with contains mucus.) intestine. GB. D. The process begins in the is broken up by chewing and is mixed a secretion from the salivary glands. bile duct. SF. The more important aspects of human digestion will serve to illustrate the way in which digestion is carried on in the higher animals. extends downward. gall bladder. end of esophagus. BD. P. C. near it and are connected with the mouth by ducts. rectum. ascending colon. Wieman. . Saliva mouth. and finally. VA.

of the proteins. This is termed peristalsis. trypsin. There is also little or no diges- tion of fats in the stomach. of the alimentary canal. changing them to peptones and The sole function of rennin is to coagulate milk and thus to separate from it the protein casein. the time it remains there depending upon both the nature and quantity of the food eaten. malt sugar. Normally the stomach becomes empty within 4 hours after an ordinary meal has been eaten. and partially digested. reduced to a semifluid consistency. (or lipase). There is no digestion of starch in the stomach except from the saliva mixed with the food. or partially digested food. little digestion The principal value of chewing is to reduce the size of the pieces of food so that fluids in the stomach and intestine can reach them quickly. FUNDAMENTALS OF BIOLOGY which acts upon starch.4 per cent hydrochloric acid and two a type of movement displayed also by other parts The food now enters the stomach. In the act of swallowing. it merely serves to activate the gastric they can do their work only in an acid medium. as Pepsin acts upon some proteoses. at first in smaller. where it is thoroughly mixed. and steapsin fatty acids. the chief ones being in the pancreatic fluid. Upon entering the small intestine. and proteoses to amino acids. the food passes down the esophagus by waves of muscular contraction in its wall. especially in the first food to enter an empty stomach. These are three in number: amylopsin. starch digestion soon ceases in the presence of the hydrochloric acid in the gastric juice. it passes by peristaltic waves into the small intestine. temporarily stored. the pyloric valve.204 called ptyalin. time. which converts starch to malt sugar. peptones. which changes proteins. as ptyalin acts only in a neutral or an alkaline medium. situated at the junction of the stomach and intestine. The in digestion. is acted upon by other enzymes. the chyme. enzymes pepsin and rennin. which breaks down fats into glycerin and . gradually converting it to Because food remains in the mouth only a short occurs there. acid plays no direct part enzymes. Innumerable glands in the wall of the stomach produce a secretion called gastric juice. After the food taken into the stomach has reached the proper degree of fluidity and digestion. This movement is controlled by a large circular muscle. but later in larger amounts. that contains about 0. However.

called erepsin. nearly all the work of digestion is in the small intestine. very little going on in the stomach or in the large intestine. maltase acts on malt sugar. Bile. Toxic products of decomposition may also be absorbed to some extent if the elimination of fecal matter from the body is considerably delayed. finger-like projections called villi. it is also acted upon by several enzymes contained in the intestinal juice. Absorption. 144). normally stays in the large intestine about 24 hours. but its presence makes conditions favorable for the action of the pancreatic fluid. especially in its upper The remaining end.METABOLISM IN ANIMALS 205 As the food leaves the stomach it is acid but is soon neutralized the pancreatic fluid and bile. Three others change complex sugars Of these. which greatly increase its absorbing Each villus contains a network of blood capillaries surface. As the food passes through especially in its digestion of fats. chiefly sugars arising from the breaking down of carbohydrates not previously digested (such as cellulose). by the small intestine. brought about by peristalsis. Some of the decomposition products. before passing out of the body as feces. on cane sugar. This is necessary because the pancreatic and intestinal enzymes act only in a neutral or a slightly alkaline medium. The absorption of food takes place almost entirely in the small intestine. time it loses most of its water and undergoes partial decomposition through the activity of bacteria that flourish in the large intestine in enormous numbers. and lactase on milk sugar. surrounding a central lacteal. The inner wall of the small intestine is infolded and covered with innumerable small. portion of the food. consisting chiefly of indigestible residue. contains no enzymes. may be absorbed here. the secretion from the liver. or slightly longer. The passage of food through the small intestine. One ments the action of these enzymes. On the outside is a layer of epithelium containing scattered mucous cells. a fluid secreted by glands in the intestinal wall. or lymph capillary (Fig. sucrasc to simple sugars. Although this is approximately the same length of time that the food may remain in the stomach. both of which are alkaline. The digested foods pass into the villi by diffusion . consumes about 4 hours. or often carried on here The During this longer. suppleof trypsin in the conversion of peptones and proteoses to ammo acids. large intestine contains no enzymes.

often interfering with their normal functions. is habitually eaten. fats supply most of the energy required by the animal body. changed back to sugar and gen. VII). liver. skin and around the heart. which Here much goes to the liver. transported. The products fats. The final step in constructive metabolism is assimilation fats is Food the transformation of digested food into protoplasm. is Chemically. glycogeu Artery Vein Diagram of an villus of man. in fact it " often referred to as animal starch. especially of and carbohydrates. to the tissues.206 FUNDAMENTALS OF BIOLOGY arising from the digestion of carbohydrates and proteins enter the blood stream directly. but carbohydrates of energy. as needed. kidneys. used both in the building up of protoplasm and as a source Proteins may serve either purpose. taken up by the circulatory system. of the sugar transported by the blood ble is converted to an insolu- Epithelium carbohydrate called glycoand temporarily stored in Later it is gradually this form. The breaking down of organic matter in the Respiration." Fi 144. some of it may be stored permanently in fatty This accumulates in thick layers beneath the or adipose tissue. and are especially important as sources of heat. but the fat derivatives There they are recornbined into are absorbed by the lymph. is eventually carried to all parts of the body to be assimilated by the living tissues. body to liberate energy has been adequately discussed in connection with plants and need not be repeated here (see Chap. When an excess of food. and intestines. is similar to starch. The digested food. intestinal Some glycogen is also stored in the muscles. which empties into a vein in the upper thoracic region. eventually reaching the blood stream by way of the thoracic duct. They In animals this energy is expended chiefly as mechanical and heat . The veins Blood capif/ctr/es carrying food from the digestive Mucous cell Lacteal organs come together to form the large portal vein.

while carbon dioxide passes in the opposite The greater structural complexity of the earthworm direction.") Fro. The a insects carry on respiration Their in unique manner. Honeybee. 145). Gills ar^ lusks. oxygen is absorbed through the surface of the body from the surrounding medium and slowly diffuses inward. flatworms. It has 207 been seen that respiration is essentially the same always involving the absorption of cells the liberation of carbon dioxide.METABOLISM IN ANIMALS onergy. consisting in most cases of either tracheae. membranes commonly occurring air-breathing in sheets or in Lungs occur in vertebrates. or lungs. Air sphere by means of small openings in the body wall called spiracles (Fig. present in most of the aquatic annelids. moland arthropods. Instead. hand. In the higher animals a respiratory system becomes established. highly branched tubes that communicate with the atmocarries food 110 dioxide. (From "Anatomy of the 8nodgrass. 145. but is blood oxygen or carbon brought directly to the cells through a system of tracheae delicate. the gaseous exchange necessary to respiration occurs in either gills all Respiratory systhe honeybee. air does not come in direct contact with the tissues. tem of parts of the or lungs. necessitates the development of a circulatory system for the transportation of the respiratory gases. ion the other In most of the higher animals. process in all organisms. and In oxygen by living in and such metazoans as sponges. and roundworms. as well as in the fishes and amphibians (at least in the younger stages of the all latter). They are spongy organs containing numerous small cavities into which air passes . aeration taking place through the moist outer skin. oxygen being transported from these organs to body by the circulatory system. coelenthe protozoans simple terates. but no special respiratory organs are present. gills. arising embryonically as pouches from the pharynx. echinoderms. They consist of thin delicate tufts.

208 directly FUNDAMENTALS OF BIOLOGY from the atmosphere. forced into and out of the lungs by movements of the diaphragm and chest muscles. from the abdominal cavity. about by relaxation of the diaphragm and chest muscles. in Decrease in the size of the thoracic cavity. the glottis So that food will not get into the respiratory is provided with a lid. and it is through exchange of respiratory gases takes place. surrounded called alveoli or air sacs. contained in the lungs. goes through the nasal passages to the pharynx. containing the vocal cords. the gas exchange that goes on in the tissues. Only about one-sixth of the air in the lungs is renewed with each breath taken. is The trachea. A distinction is often made between external respiration. while in internal respiration it receives carbon dioxide and liberates is inhaled through the nostrils. The larynx. The absorption of oxygen by the blood and the liberation of carbon dioxide take place by diffusion through the delicate walls of the capillaries and of the alveoli. Both gills and lungs are richly their walls that the supplied with capillaries. the ribs and sternum are pulled outward by the chest muscles. by capillaries. Thousands of these alveoli. a tube that lies in front of the esophagus it and passes downward into the thoracic cavity where In the lungs these break tubes. which the lungs lie. thus forcing air out of them. a" feature of all Air is is a muscular partition separating the thoracic cavity. which closes during the act of swallowing. When relaxed. In man. forms the upper part of the trachea. and air rushes into the lungs. diaphragm contracts. brought to expand. gives rise to up a pair of bronchial into smaller and smaller branches. or windpipe. thence into the trachea. The diaphragm. In external respiration the blood receives oxygen and liberates carbon dioxide. The opening from the pharynx into the larynx is termed the tract. and finally through the bronchial tubes into the lungs. and internal respiration. Under ordinary conditions about a pint of air is inhaled and exhaled. glottis. the exchange of gases occurring in the gills or lungs. causing them mammals. As the it becomes flat and thus enlarges the thoracic cavity. approximately two and a half quarts . the diaphragm is arched upward. air voice box. are . or oxygen. the epiglottis. which finally terminate in minute thin-walled cavities. but when contracted. makes possible the contraction of the lungs.

oxygen. . when the breathing is deeper and more rapid. but in these groups the blood flows from the open ends of vessels into sinuses. in the higher animals. and metabolic wastes takes place through Blood is propelled through its course by the walls of capillaries. The color of blood due to the presence of a pigment present either in the plasma. the air exhaled from the lungs contains about 16 per cent of oxygen and 4 per cent of carbon dioxide. and waste materials is Thus in the roundworms and echinoderms a colorless required. a special means of transporting digested food.METABOLISM IN ANIMALS 209 remaining in the lungs. Cells not in direct contact with food or oxygen circulatory fluid. Likewise in the mollusks and arthropods a definite vascular system occurs. red blood to hemoglobin. oxygen. are close to cells that are and can readily absorb from them these vital necessities. have a closed system of circulation. so called because it performs the dual functions of digestion and circulation. blood owes its color to hemocyanin. Since the atmosphere contains approximately 20 per cent of oxygen. the blood remaining in the vessels throughout its course. are of two kinds: red corpuscles and white corpuscles. Similarly. of its oxygen and takes up about an equal amount of carbon dioxide. or. The blood cells of vertebrates. The blood is of insects is colorless. while the blood of other is all vertebrates red. circulatory fluid containing amoeboid cells is present in the coelom. In animals of greater structural complexity. The lower animals have no need for a special Circulation. annelids. carries digested food and oxygen directly to all parts of the body. suspended in the colorless plasma. while in the annelids. waste materials have to move only a In the coelenteratos and flatshort distance to be eliminated. one or more contractile vessels heart. as in vertebrates. however. worms the gastrovascular cavity. Blue as in invertebrates. only about a quart The inhaled air loses about one-fifth of air is left in the lungs. a closed system of vessels through which blood circulates is developed. in addition to the coelomic fluid. or in the corpuscles. like the is designated as an open system. During violent exertion. by a cells Blood. Such a circulatory system The vertebrates. Here the transfer of digested food. that of many crustaceans and mollusks invertebrates and of is blue. Blood consists of a liquid plasma containing called corpuscles.

The corpuscles and the plasma have different functions. after up its oxygen to the becomes hemoglobin Because the color of again.000 to the When oxygen is taken up by the red corpuscles. stable ^10 giving tissues. In shown.000 to 9. compound that. hormones. The two meter. moving by pseudopodia. pure (aerated) blood is X 1. a substance that aids in the clotting of the blood. 146). largest kinds of white corpuscles exceed the red* blood cells in size and may become amoeboid. 146.000. two of them are shown in edge view. it combines loosely with other v }\ / /^ A : fc / r^\ G') ^^ i^^ /&\ . human cubic millimeter. and fibrinogen. to exceed 4 weeks. it. The red corpuscles lack nuclei. which is This is an unbrilliant red. A peculiar feature of white corpuscles is their ability to pass .000 to the cubic milli- Three principal kinds occur. the red corpuscles lose their nuclei soon after being all other vertebrates they remain nucleated (Figs. not Hence new ones are constantly being formed. of corpuscles are especially concerned with the transoxygen. Human blood corpuscles.:A ^ark-red hemoglobin to form oxyhemoglobin. in corpuscles are much and liver. which differ in size and in the character of their nuclei. disk-like in man and In (Fig. in form. It consists of about 90 per cent water. blood depends upon the presence or absence of oxygen in FIG. Dead corpuscles origin being red bone marrow. They are oval or circular. The red portation cells. mammals formed. mammals being biconcave disks blood they number about 5. abundant than the red man numbering from 5. Three white corpuscles are bright red. while impure (non-aerated) blood is dark red or slightly purplish. their place of are destroyed in great Red blood cells do not live long in man. numbers in the spleen less The white ones. The plasma carries most of the dissolved food to the tissues and removes carbon dioxide and other waste products from them. antibodies. In one kind the nucleus consists of several lobes with delicate connections between them.210 FUNDAMENTALS OF BIOLOGY Only the former contain hemoglobin. and also contains inorganic salts. but in 141 and 146).000.

severe infection. empties into a vein in the upper thoiaoic region. They lack Their nature. of It known as lymphatics. which are constituents only of mammalian blood. and wornout cells. bring about the This is circulation of accomplished in the body. water. vertebrates have a colorless It occupies intercellular fluid derived from it called lymph. In addition to the blood. in closing the fluid. Its function is thus supplementary to that of the blood. Their function consists partly in engulfing small solid particles.METABOLISM IN ANIMALS 211 through the walls of capillaries into the surrounding tissues. which is now designated as serum. further loss of wound. origin. In certain types of attracted to places of injury or infection. which are contractile organs present in the lower vertebrates but not in mammals. White corpuscles are formed nodes. the living cells of the body and acts as an intermediate. prevents Lymph. waste material. the number of white blood cells increases two or three times. as previously noted. . They seem to assist in the coagulation of the blood. cles. the largest lymphatic. They also aid in the transportation White corpuscles are of fats to different parts of the body. blood. Lymph consists mainly of plasma and white corpuscles that have escaped from the blood Lymph surrounds vessels. In so doing. Lymph hearts. Lymph nodes are enlargements of lymphatics where white corpuscles are formed and where bacteria that have entered the body are destroyed. and in the removal of waste products. are smaller than red corpuscles and colorless. functions are not well understood. This occurs when blood comes in contact with air or with injured tissue. lymph throughout of the muscles. it squeezes out the remaining The clot. and nuclei and are not regarded as cells. in red bone marrow and in lymph Blood platelets. such as bacteria. which they destroy by digesting them. medium 1:1 the transportation of nourishment and oxygen to them from the blood. lymph spaces and vessels circulates throughout the body in delicate mals. which are best developed in mamMost of these eventually connect with the closed system blood vessels through the thoracic duct. The fibrinogen of the plasma is converted to threads of fibrin that entangle the corpus- forming a clot. from contraction mammals by pressure resulting Excretion. The breaking down of organic matter in the the animal body results in the formation of carbon dioxide.

the skin. passes out of the body through the and kidneys.212 salts. 147. In the vertebrates urea is formed chiefly in the liver and is then carried by the blood to the kidneys. FUNDAMENTALS OF BIOLOGY and other metabolic waste products. and the excretory organs. and other waste This passes materials. Nitrogenous waste products. where it accumulates. is dissolved in water to form urine. The urea. always result from the decomposition of proteins and amino acids. These are collected by the blood stream and eliminated through the respiratory In mammals nearly organs. from the kidneys through the ureters to the bladder. all the carbon dioxide is given off through the lungs. salts. principally urea. with a smaller quantity of uric acid. Diagram of human excretory system. while water lungs. Kidney B/aMer Ure -f~hrcr Fia. one of the kidneys shown hi longitudinal section. skin. where it .



stored before passing out of the

body through the urethra

(Fig. 147).

the kidneys are a pair of bean-shaped bodies situated in the posterior part of the abdominal cavity close to the dorsal body wall. They are composed mainly of a large number of



slender complexly branched tubules, each having a terminal Water with its capsule that surrounds a network of capillaries.

waste substances passes by diffusion from the capillaries into the



Sweat duct


muscle ofthe hair



Sweat gland Blood

FIG. 148.


Somewhat diagrammatic section of the mammalian Wolcott, "Animal Biology.")


renal tubules.


lead to larger and larger tubes, into the ureter. finally converge The human skin comprises an outer portion, the epidermis, and an inner portion, the dermis (Fig. 148). The epidermis is

The tubules


of stratified

epithelium, the dermis of connective

The dermis

also contains sensory nerve endings,


ous blood vessels, sweat glands, hair follicles, sebaceous glands, and fat cells. Excretion occurs mainly through the sweat glands, each of which is a slender tube opening to the surface by means The lower end of the sweat gland is highly coiled and of a pore.

surrounded by a network of capillaries from which water and small quantities of waste products pass. The evaporation of



perspiration leaves a residue of salts and other waste matter on the skin. Sweat glands are stimulated by high temperature,

the increased


of evaporation tending to keep the


hair arises at the


of a hair follicle, a

body narrow


pocket formed by the infolding of the epidermis into the dermis. follicle is surrounded by sebaceous glands, which secrete

secretion rather than an excretion.

This softens the hair and skin, and so may be considered a Sebaceous glands are present
parts of the



body except the palms

of the

hands and the

soles of the feet.



Irritability, or the capacity to react to external influences, is a fundamental property of protoplasm and therefore is universal among living things. Although response to stimuli is no more definite in animals than in plants, almost always it is far more rapid and is brought about by a very different mechanism, viz., In the chiefly through the cooperation of nerves and muscles. of cells the that the lowest react to group sponges, metazoans, external influences are stimulated directly, there being no nerve cells. In all the higher animal groups, however, this is not the Here differentiated cells are present whose sole function case. is to receive and conduct stimuli and to bring about responses in other tissues that, for the most part, are themselves incapable In the coelenterates, simple nerve of receiving stimuli directly. cells are present but are not aggregated to form ganglia. In the higher groups, the nervous tissue is organized as an elaborate system of nerves and ganglia, different types of which have already been considered in a number of animals. The nervous system, in controlling the responses of an animal to external influences, makes it possible for various groups of cells to act In other words, it directs and coordinates vital funcas a unit.




controlling influence, there could be


bodily activities.

Nervous System
general plan in



Although constructed on the same

vertebrates, the nervous system reaches its highest degree of organization in man, the brain, in particular, greatly surpassing in relative size and complexity that of any other* animal. The central nervous system comprises the brain



spinal cord, the peripheral nervous system all the nerves from them. The brain consists of three main divisions:

The cerebrum

the cerebrum, the cerebellum, and the medulla oblongata (Fig. 149). is by far the largest division. Its cortex, or outer


of gray matter, its inner portion of white 215


The amount

greatly increased by the development of convolutions, or furrows, which are deeper and more numerous in man than in any other mammal. The cere-

gray matter


bellum, lying beneath and behind the cerebrum, is much smaller and only slightly convoluted. The medulla oblongata is a bulbous enlargement situated at the upper end of the spinal cord.

Lateral f/ssure

Centra] sulcus


Fia. 149. Side view of the human brain. The roots of the cranial nerves are indicated by roman numerals. (From Wolcott, "Animal Biology.")

In man, 12 pairs of cranial nerves arise from the brain. Most of these extend to the sense organs and to other parts of the head, but some go to the neck, shoulders, and organs of respiration,

and digestion. It is almost spinal cord lies inside the spinal column. circular in cross section and is nearly divided into symmetrical


fissures, one dorsal and the other ventral In the center is a very small canal. This is surrounded by gray matter, which extends into each half, through


by two deep

(Fig. 151).




the outer mass of white matter, to form a dorsal and a ventral As seen in cross section, the gray matter has somewhat

the form of the letter H. The spinal nerves, of which there are 31 pairs, arise from the spinal cord and pass outward between the vertebrae to supply the lower portions of the body. The cerebrum is the seat of intelligence, consciousness, memory,

and the higher emotions.

It controls all voluntary actions. The cerebellum coordinates muscular activity, particularly the movements concerned with equilibrium and locomotion. The medulla

oblongata governs such reflex actions as winking, sneezing, and It regulates the activities of the organs of respiration coughing.


of the alimentary tract

movements and the secpetion of digestive juices. The spinal cord transmits impulses to and from the brain and is the center of reflex actions involving the trunk and limbs.
of circulation.
It also controls the peristaltic

The sympathetic nervous system comprises a number of nerves, ganglia, and plexuses, or groups of ganglia, situated in various parts of the body outside the central and peripheral nervous chain of these ganglia lies on each side of the spinal systems. cord and is connected with branches of the spinal nerves. The


regulates such involuntary activities as the beating of the heart, the secretion of glands, and the contraction of all involuntary muscles.

cardiac plexus the stomach.


just below the heart, the solar plexus behind

The sympathetic nervous system

Neurons and Nerves. The neuron, or nerve cell, is the strucand functional unit of the nervous system. It consists of a central nucleated portion, the cell body, and a number of extremely
fine cytoplasmic extensions called nerve fibers, which conduct nervous impulses (Fig. 138). The fibers of each neuron are of two kinds, distinguished from each other by the direction in which the impulses are normally conducted by them. Dendrites carry impulses toward the cell body, while axons carry them away from the cell body. Commonly a neuron possesses a number of relatively short, branched dendrites and a single long axon usually branched only at its tip. Generally the dendrites and the axon extend in opposite directions. It should be understood that a nerve cell, as a whole, ordinarily can transmit impulses in only one direction. In going from one nerve cell to another, the impulse passes froin the terminal branches of the axon of the one neuron to the dendrites of the



The fibers of the two neurons are close together but probably not in organic union, the connection between them being termed a synapse. A sensory neuron carries impulses from a point of stimulation to a nerve center, such as the brain or spinal cord, while a motor neuron transmits impulses from a norve center to a tissue in which a response is to be induced; such as a muscle or a gland. The structure stimulated is called a receptor the one that responds, an effector. In vertebrates, nearly all the cell bodies are in the gray matter of the brain and spinal cord, the others occurring in the

ventral ganglion



.sensory nerve fiber

motor nerve


longitudinal muscles''



circular muscles FIG. 150. Diagrammatic cross section of the ventral nerve cord and surrounding structures of an earthworm. (From Parker, in Popular Science Monthly,
after Retains.)

ganglia of the peripheral and sympathetic nervous systems. The white matter of the central nervous system consists chiefly of

Ganglia are merely aggregations of cell bodies, while nerves are bundles of nerve fibers usually enclosed by a sheath of connective tissue. Thus a nerve fiber is like a wire, a nerve like a cable. A sensory nerve consists of the fibers


(dendrites) of many sensory neurons, a motor nerve of the fibers Mixed nerves, on the other (axons) of many motor neurons.

hand, are composed of both kinds of fibers, and so are both sensory and motor in function. Reflex Action, A reflex is a movement or other act resulting from the transmission of an impulse inward from a receptor to a



nerve center and then outward to an effector. It is the simplest type of nervous action. The impulse concerned travels along a path designated as a reflex arc. This always involves at least one sensory and one motor neuron. A simple illustration of

may be seen by touching the skin of an earthworm. The mechanism by which a response to this stimulus is brought about may be understood by studying Fig. 150 in connection
reflex action

with the following account. Lying in the skin of the earthworm are a number of sensory neurons, the dendrites of which receive stimuli acting upon the surface of the body. The axon of each of these neurons passes inward to one of the ganglia of the ventral nerve cord, where its end comes into synapse with the dendrites of one of the many motor neurons of which the ganglion is composed. The axons of certain of these motor neurons pass outward to the muscles of the body wall. Stimulation of the skin causes an impulse to be sent over one or more sensory nerve fibers to the central nervous system. There it is transmitted by one or more motor nerve fibers to a group of muscles that is thereby stimulated to contract. This behavior is called reflex action because the impulse is reflected from the nerve center somewhat as light is reflected by
a mirror. Intermediate neurons, connecting the sensory neurons stimulated with other motor neurons, may transmit the stimulus
to other muscle cells and, as a result of their contraction, the

earthworm may crawl away.

Most of the bodily activities


man are brought about by reflex

action, and, in such cases, a response to a stimulus is induced without volition. For example, if one's hand accidentally

touches a hot object, it is quickly withdrawn before the sensation of pain reaches the brain, because the motor impulse is sent back to the muscles of the arm directly from the spinal cord. The
constriction of the pupil of the eye in bright light and its dilation in partial or complete darkness are involuntary responses brought about through reflex action. The acts of sneezing and coughing


are reflexes, as are also the secretion of gastric juice, the peristaltic of the digestive tract, changes in the rate of beating

of the heart, the

movements concerned with breathing, shivering

In breathing and similar acts, the stimulus cold, etc. involved arises from within the body instead, as in other reflexes,


coming from outside.



The Spinal Nerves. In vertebrates, the neurons concerned with most of the reflexes occur in the spinal cord. Sensory nerve fibers lead to it and motor nerve fibers lead from it to the muscles. Each spinal nerve arises from the spinal cord by means of a dorsal and a ventral root (Fig. 151). These spring from the horns of gray matter in the cord. The dorsal root consists of sensory nerve fibers, the ventral root of motor nerve fibers. The cell bodies of all sensory neurons entering the spinal cord lie in a ganglion that forms a slight swelling near the base of the dorsal

The ventral




without a ganglion,


made up

Gray matter

Dorset/ root


Effee -for Fio. 151. Diagrammatic cross section of the spinal cord with dorsal and ventral roots of a spinal nerve, showing neurons involved in a simple reflex arc. Arrows indicate path of a nervous impulse.
of the axons of

Motor neuron Wh/te matter Ventrafroot

motor neurons whose


bodies and dendrites

in the gray matter of the spinal cord. Some of the fibers of the dorsal root are directly in synaptic contact with the dendritas

of the

motor neurons

in the spinal cord, while the connection of


through one or more intermediate neurons.

nerve, after arising by three branches, each of which contains both sensory and fibers. These branches go to different parts of the body.


and ventral

Each spinal divides to form root,

In vertebrates, each neuron is connected with many other neurons, so that impulses may be transmitted along many paths. This makes possible different types of responses as well as the coordination of activities. Thus a stimulus causing a reflex action may reach the brain, and the response may be modified or supplemented by voluntary actions involving other parts of
the body.



Sense Organs. A sense organ is an organ specialized to receive a particular kind of stimulus, transform it into a nervous impulse, and transmit it to the brain. The organs of special sense are those concerned with sight, hearing, smell, taste, and touch. Receptors sensitive to touch are distributed over the entire
surface of the



body but are especially numerous on the palms The sense of taste is localized in groups of sensory



particularly on the tongue,

These are present in the mouth, and are connected with branches of


FIG. 152.


of the



(After Czermak.)

the seventh and ninth pair of cranial nerves, by taste sensations are transmitted to the brain.

means of which The taste buds

can be stimulated only by substances in solution. Moreover, only four kinds of tastes can be distinguished sweet, sour, bitter, and salty. Other apparent tastes are due to volatile substances
that affect the nerves in the nose, and so are really odors. The sense of smell is localized in the mucous membrane of the nasal

This contains sensory cells connected with nerve cavities. endings of the olfactory nerves, one of which leads from each Smell is a far more delicate sense than taste nostril to the brain.




better developed in such animals as the dog than




called the malleus. The Eye. and white. The ear is an organ specialized for the reception of sound. is kept moist and free from dust particles by a secretion from the lachrymal gland. The eyeball is moved by six Its surface muscles. connecting the tympanic membrane with the inner car. and the inner ear (Fig. It comprises a vestibule into which open three semicircular canals and the The semicircular canals are concerned spirally coiled cochlea. which serves to equalize the air pressure on either side of the eardrum. extends across the middle ear. These set up vibrations in the eardrum. opaque. is dense. the On the inner side of the eardrum a small air-filled cavity. or outer coat. 152).222 FUNDAMENTALS OF BIOLOGY The Ear. and is protected externally by an upper and a lower eyelid. or membranous labyrinth. Over the surface of the membrane that lines the cochlea are numerous sensory cells in contact with delicate terminal branches The outer ear acts as a trumpet. The coat. ing sound waves and directing them into the ear canal. At the front of the eye it becomes the transparent cornea. from which numerous ducts lead to the inner surface of the upper eyelid. the three small bones of the middle ear to the liquid of the inner The sensory cells in the lining of the cochlea transform the sound vibrations to nervous impulses that are transmitted to the brain 153. In man and etir. A chain of three small bones. outer ear leads to a canal. surface of the eyelids. incus. not with hearing but with equilibrium. middle ear. is a closed. the sense of hearing depending upon the transfer of impulses by the pair of auditory nerves from the ears to the brain. the inner which is closed by the eardrum. a thin transparent membrane continuous with the mucous membrane that lines the inner The sclerotic. is It has three coats: the sclerotic eyeball is nearly spherical. by the auditory nerve. Both the sclerotic coat and the cornea are . This is connected with the throat by the Eustachian tube. fluid-filled sac. The structure of the human eye It lies in is shown by Fig. is or tympanic membrane. other mammals the ear consists of three parts: the outer The end of the middle ear. collectof the auditory nerve. an orbit. which are then conducted by ear. or eye socket. and the retina. The inner ear. the choroid coat. and stapes. about 1 inch in length. The front of the eyeball covered with the conjunctiva. four of which are straight and two oblique.

i. v. ret. Diagrammatic longitudinal section of the human eye. The focus of this lens system is controlled by the sus-. aqueous humor. are called rods and cones. ciliary muscle. the vitreous humor. pupil. ul. suspensory ligament. scl. forming a thin lining inside the choroid coat and extending forward almost to the iris. opt n. cornea. conj. or middle coat. sclerotic coat. retina. In iris is an opening called the pupil.COORDINATION IN ANIMALS thick and tough. p. oil. Both the choroid coat and the iris are provided with a pigment that prevents light from entering except through the cornea. upper lid. crystalline lens. crystalline lens. aqueous humor. I. It is very complicated in structure. and the cornea is filled with a transparent liquid. sus. while the much larger space behind the lens contains a transparent gelatinous substance. The retina is the inner coat of the eyeball. chj choroid coat. . The expansion and contraction of the iris change the size of the pupil and regulate the amount of light passing through the lens. II. cor. vitreous humor. the aqueous humor. lower lid. The The space between the lens lens is biconvex and transparent. These Behind the iris is the crystalline lens. The cornea. the center of the op-f n FIG. iris. attached to the choroid coat by the suspensory ligament. a. optic nerve. consisting of two kinds of neurons specialized for the reception of light. 153. ous blood vessels. of the cornea 223 The choroid. and vitreous humor constitute a complex lens system that throws an image upon the retina. forming the iris. conjunctiva. contains It separates from the sclerotic coat just numerback and hangs down as a curtain.

It consists of two lobes. unlike the products of other glands. produce nervous impulses that are transmitted by the optic nerve of the made up to the brain. hormones regulate and coordinate certain functions. developing into an undersized frog. Its chemical formula is CisHnC^NI^ Thyroxin con- trols the rate of oxidation in the body and so has a profound . For example. Thyroid Gland. Light stimuli. and the pituitary body. examples being the pancreas and the glands. These arc secreted by glands but. a tadpole fed on fails to occur.224 FUNDAMENTALS OF BIOLOGY pensory ligament. on the other hand. attached to which are delicate ciliary muscles. certain functions are controlled by special substances called hormones. or ductless sexual organs (testes and ovaries). the development of a tadpole into a frog is determined by a hormone produced by the thyroid gland. thymus. These regulate the tension of the suspensory ligament. In general. which are connected by intermediate neurons with the lightwensitive rods and cones. Although the coordination of bodily brought about mainly through the agency of the nervous system. a substance that has been found to contain about 65 per cent of iodine. It penetrates the sclerotic It is and choroid coats at the back of the eyeball. The optic nerve connects the retina with the brain. from a deficient or an excess secretion of any hormone. one lying on either side of the trachea just below the larynx. when focused on distant objects it is made less convex by relaxation of the ciliary muscles. growth continues but metaOn the other hand. generally acting as either activating or Nearly always an abnormal condition arises inhibiting agents. arc not carried by ducts but pas directly into the blood or lymph. These include the thyroid. If this gland is morphosis removed from a young tadpole. adrenals. thyroid extract undergoes a premature metamorphosis. parathyroids. other functions as well. axons of sensory neurons. are special hormone-secreting organs. activities is Chemical Coordination. lying in the retina. The endocrine. which in turn changes the thickness and curvature of the crystalline lens. falling on the retina. this process being known Some organs that produce hormones have as internal secretion. When the lens is focused on nearby objects it is made more convex by contraction of the ciliary muscles. This gland secretes thyroxin. The thyroid gland is situated in the front of the neck.

Exophthalmic goiter is due to an excess secretion of thyroxin. two on either side. prepared from the thyroid glands of cattle or of other animals. or outer covering. usually arising from a degeneration of the gland itself. either during childhood life. consists of a cortex. etc. It is usually < . from an insufficient amount of iodine in the diet. protruding eyeballs. characterized by a low rate of metabolism. this usually resulting.COORDINATION IN ANIMALS influence 225 on physical growth and mental development. a peculiar thickening of the skin. Degeneration of the thyroid during adult life produces a disease called myxedema. They secrete a h^mone that regulates the amount of calcium in the blood. The cortex secretes cortin. by surgical Thymus Gland. The thymus. etc. Removal of the parathyroids results in violent muscular contractions followed by death. These two small bodies. arising and physical and mental sluggishness. excessive weakness. secreted also Parathyroid Glands. artior of thyroid extract. This disease is characterized by a high rate of metabolism. one lying above each kidney. deficiency of thyroxin during childhood. Adrenal Glands. Its functions are imperfectly known. and a medulla. results in a form of dwarfiiess A and idiocy known as cretinism. It is conspicuous in childhood but gradually becomes smaller during adolescence. The symptoms from underactivity of the thyroid. removal of a portion of the thyroid gland. A deficiency of cortin produces Addison's disease. by the eating of sea food or by the use of iodized table salt. a small bilobed gland. in turn. lies in the upper part of the thoracic cavity below the thyroid. The hor- mone and by the thymus gland has an effect upon growth seems to be responsible for the arrested development of the sexual organs during childhood. An excess secretion results in an increase of calcium accompanied by softening of the bones and by muscular weakness. emaciated condition accompanied by a darkening of the skin. or adult may ficially synthesized. nervousIt may be relieved ness. in most cases. Goiter is is an enlargement of the thyroid gland. be corrected by the feeding of thyroxin. or epinephrin. or central portion. It may be remedied or prevented. The parathyroids are four small glands attached to the dorsal surface of the thyroid. Simple goiter caused by a subnormal secretion of the thyroid hormone. the medulla adrenalin. an anemic.

and the pancreas. an excess production about of which. to arrest hemorrhage. causes contraction of the blood and to the can added muscles. a hormone secreted into the blood by certain groups It regulates the normal metabolism of carbohydrates. but which have other functions as well. and feet enlarge abnormally. called tethcMn. Its injection into the blood retards the pulse. etc. stimulates the flow of their fluids. lobe of the pituitary body secretes pituitrin. the small Hormones secreted into the blood intestine. This is poured into the blood by the mucous membrane of the small intestine when the acidified chyme of the stomach comes in contact with it. by increasing the formation of glycogen Insulin is of cells in the pancreas.226 fatal. especially the size of the skeleton. certain groups of cells in the testes and ovaries are responsible for many of the bodily changes that occur during adolescence. gives rise to gigantism. stimulates the heart and brain. upon reaching the liver and pancreas. bodily growth. and by decreasing the amount of sugar produced from . and causes contraction of the larly those of the intestine and uterus. . hands. secretion of this An regulates over- hormone during youth in while an undcrsecretion results dwarf ness. a disease in which the bones The posterior hormone that of the face. brought by the emotional states of anger or fear. The anterior lobe secretes several at the base of the brain. Adrenalin is used in medical practice to stimulate the heart. Adrenalin be strength prepared gives from the adrenal medulla in purified crystalline form and can Its chemical formula is C 9 HuNO 3 also be made synthetically. by For example. vessels. increases the concentration of sugar in the blood. Other Glands. FUNDAMENTALS OF BIOLOGY Adrenalin is a powerful stimulant. The secretin. raises the blood visceral muscles. different hormones. oval. a causes smooth (involuntary) muscle to contract. This is a small. It does this by accelerating the rate of sugar oxidation in the tissues. bilobed body occurring Pituitary Gland. in the liver. An oversecretion of tethelin during adult life produces acromegaly. particu- pressure. include the sexual organs. The production of bile and pancreatic fluid i^brought about by a hormone called secretin. a hormone produced by the testes brings about the development of the beard and deeper voice. to raise the blood pressure. One of these. Organs producing hormones.

and permits the .COORDINATION IN ANIMALS fats 227 and proteins. Diabetes is a disease resulting from an insuffiIts symptoms are cient secretion of insulin by the pancreas. Diabetes may be controlled by a restricted use of carbohydrates in the diet and by the injection into the body of insulin prepared from the pancreatic glands it of sheep or cattle. This treatment does not cure the disease but merely relieves the symptoms arising from patient to lead a more healthy life. mainly an increase in the amount of sugar in the blood and the presence of sugar in the urine.

accompanied by more tion. while in asexual reproduction no such fusion occurs. however. in Planaria. occurring only in certain protozoans. or less cell differentiaincell In unicellular organisms. cell on the other hand. resulting in reproducIn animals. growth volves only division tion. Asexual Reproduction.) 1 is far more prevalent than the latter. have seen that among the proto- up an independent existence. in Hydra. but merely an In all multicellular increase in size of one already in existence. although characteristic of nearly all plants. 154. Occasionally this happens among some of the lower metazoans. is very rare among animals. after Child. the proby which an organism gives rise to others of its own kind. except among the lowSexual reproduction est groups. We zoans the prevailing method of reproduction is fission an equal division of the body into two new parts that separate and take always involves a fusion of two gametes. 326-328). Spore reproduction. (From A. reproduction may be either sexual or asexual. Wieman. A. "General Zoology. although the former Fission in metazoans. as in Hydra and the flatworm Planaria (Fig. organisms plished cell growth is by both cell division accomand enlargement. which does not involve the production of a new individual.' after Koelitz. such as the malarial parasite (see pp. B. FIG.CHAPTER XIV REPRODUCTION AND DEVELOPMENT IN ANIMALS cess is The continuity of life is maintained by reproduction. It closely related to growth or development. Other methods of asexual reproduction are found in animals. enlargement. 154). B. particularly among the lower groups of meta228 . as in plants.

attention has already been called to the formation of buds in Hydra (see p. somewhat uncertain. blood. 125). Even many of the protozoans exhibit a form of sexual reproduction. It is only iii a relatively few protozoans that gametes are formed and a permanent union takes place between them. an additional band of cilia occurring at the posterior end of the cell. 227). the malarial parasite. and free swimming. the two cells separate. Budding is common in the sponges. one of the conjugating individuals is much smaller than the other. lateral outgrowths from the body wall that become detached and give rise to new indi- This is a method of reproduction comparable to "vegetative propagation" among plants. In Vorticella. a behavior known as conjugation takes place for Two individuals come in con(Fig. Germ egg. nerve. but even where asexual methods occur (as in Hydra). Among the viduals. Because no new individual is strictly speaking. complex nuclear transformations occur. although the rapid increase in number of individuals is brought about chiefly by fission. and other tissues that are primarily concerned with metabolic activities with the maintenance of shall soon see. higher animals. asexual reproduction vertebrates it does not occur at all. as Cells. we . and flat worms. in most cases. sexual reproduction is also present. is rare. devoid of a stalk. coelenterates. 155). In the growth of a metazoan from the fertilized most of its cells become specialized to form muscle. sexual method. under certain conditions. tact with each other. Conjugation in Paramecium. For example. often giving rise to multicellular colonies. as previously explained. not repro- duction. Nearly all animals reproduce by the In fact. The significance of conjugation has been variously interpreted but is still FIG. In Paramecium. example. In Plasmodium. this behavior is formed. it is the only way by which new individuals arise. in fact in the Sexual Reproduction.REPRODUCTION AND DEVELOPMENT IN ANIMALS 229 zoans. for example. 155. A permanent union occurs between the conjugating individuals. and after an exchange of nuclear material. eggs and sperms are differentiated (Fig.

collectively. or." Princeton University Press. Except nr. and are comparable to what are termed " vegetative tissues" in plants. B. XV). the soraa. (From Conklin "Heredity and Environment. in such simple . these germ cells. and thereby become transformed into ripe gametes. after Retzius. after Hertwig. taking no part in general bodily functions. As the body grows. however. by permission.230 FUNDAMENTALS OF BIOLOGY the individual of which they are a part. a relatively few remain unspecialized. they undergo a period of multiplication but are incapable of functioning as reproductive cells until the animal reaches maturity. The ovum (FC) inside of which is the clear membrane (Memb) and within this is the ovum proper containing yolk granules (F) and a nucleus (N) embedded in a clear mass of cytoplasm. Then they undergo certain complex changes (described in Chap. 156. A nearly ripe human ovum surrounded by follicle cells metazoans as the hydra. Of all the cells that arise from the fertilized egg. B. -FO is in the living condition. These are called somatic cells. are set apart at a very early stage of embryonic development. two human spermatozoa drawn to about the same scale of magnification as the egg.) FIG. as they arc called. X 250.

The egg is now said to be fertilized. or shell. and some a great deal larger. The human egg cell is approximately ^25 i nc h in diameter. as among plants. 1565). and the coelenteratcs. 157).. In addition to the nucleus. The female gametes.REPRODUCTION AND DEVELOPMENT IN AWMALS 231 but in In the sponges. all other metazoans they are confined to definite sexual viz. egg. are incidental. which surof a bird's "egg. In most cases they swim by means of a flagellum or tail. 156). their eggs on land. is Thus the white portion stored in accessory layers. Among animals. The sperm swims to the egg and penetrates its outer membrane. and when the male and female nuclei fuse they may be approximately equal in size (Fig. and are nearly always non-motile (Fig. but permanent in all the higher groups. are generally spherical in shape. called eggs or ova. then gradually increases in size as it approaches the egg nucleus. Although an egg may be surrounded by millions Its nucleus of sperms. they merely aid in bringing about this gametic union or in providing a means for the new individual to develop. Fertilization. only one normally succeeds in effecting fertilization. These are temporary organs in ovaries. which constitutes the yolk. In many cases is is round the albumen. while the sperm (including the Other eggs are much smaller tail) is slightly over %QQ inch long. which lay Its chief purpose is to prevent evaporation. sperms or spermatozoa. while in others it part of the reserve food uniformly distributed throughout the accumulates at one end. are a great deal larger than the sperms but much less numerous. the latter constituting the ovum or real egg enormously enlarged by the accumulation of yolk. are The male gametes of animals." called merely accessory food material surrounding the yellow portion. a new individual arising from the zygote by repeated All other features associated with reproduction cell division. germ cells are scattered throughout the body. called of various forms but are always very small and ordinarily consist of but little more than a nucleus (Fig. than the human egg. testes organs. the essential feature of sexual reproduction is the fusion of a male and female? gamete. In some animals the yolk egg. The eggs of many animals are surrounded by a special protective envelope particularly in such forms as reptiles and birds. It is important to realize that the fertilized egg or zygote gives . all eggs have a relatively large amount of cyto- plasm containing reserve food.

an egg Parthenogenesis. both the sperms and eggs are discharged into the water in large numbers. Fertilization in one of the roundworms (Ascaris megalo- Gphald) the male and female nuclei in contact within the cytoplasm of f only in the fishes and amphibians that a similar condition exists. mainly invertebrates. the egg. and the sperms are discharged directly brates into the water. in certain small crustaceans. Here the eggs are fertilized before they are laid. and wasps (Figs. but in the vertebrates fertilization occurs in the oviducts. as in the frog. and in all reptiles. it is actually the new individual itself! In order that the sperms may reach the eggs. it is Among the verte- FIG.232 rise to all FUNDAMENTALS OF BIOLOGY the cells of the adult individual. in the starfish. polar bodies. and conse- quently to them fertilization presents no difficulties. as in most fishes. in a few fishes and amphibians. For example. there must be a liquid medium through which they can swim. for example. and in some cases there are apparently no . an unfertilized egg may give This phenomenon is known as rise to a new individual directly. The sperms are introduced into the body of the female and swim to the eggs purpose. usually over the eggs just after they have been laid. while still groups. 158 and 159). In fact. bees. the means by which the gametes are brought together must necessarily be modified. Although in the great majority of animals will not develop unless a sperm has united with it. and there fertilization takes place. or while they are leaving the body. Internal through fluids that are secreted for the fertilization occurs in the insects and in a few other invertebrate and mammals. in a group of microscopic parthenogenesis. 157. Most of the groups of invertebrates are aquatic. X 1. and in such insects as plant lice or aphids.000. aquatic forms called rotifers. The eggs pass down through the oviducts ancj out of the body before being fertilized. in some animals. Sometimes no males are produced for many generations. In some invertebrates the eggs are fertilized in the ovaries. In animals that do not live in the water. In aquatic animals this is provided by the water in which they live. It occurs. pb. birds. and among ants. where the sexes are separate.

males at all. In the bees and related insects the female lays both fertilized and unfertilized eggs, the former always develop-

FIG. 158.

Parthenogenetie animals.

A, a

rotifer; B,

an aphid;


a crustacean.


Shull, "Heredity," A, after Harring; B, after Webster; C, after StorchJ

ing into queens (fertile females) and workers the latter into drones (males).

(sterile females),

FIG. J69.






(From Phillips, U. S. Department of Agriculture
Farmers' Bulletin 447.)

Early Embryonic Stages. In the development of the embryo from the zygote, we find that the early stages are essentially similar in all metazoans, the differences that occur being mostly

in details.


This development we shall follow in a very general way, using as an illustration the lancelet (Amphioxus), a very primitive chordate previously discussed on pp. 164 and 165.



Successive stages in cleavage and gastrulation of Amphioxus. A, C and D, 4 cells; E, 8 cells; F, 16 cells; G, blastula stage of about 96 cells; H, section through same showing the cleavage cavity (blastocoele) 7, blastula seen from the left side, showing three zones of cells, viz., an upper clear 2one of ectoderm, a middle (faintly shaded) zone of mesoderm, and a lower (deeply shaded) zone of endoderm cells; /, section through same showing these three types of cells; and L, successive stages in the infolding of the endoderm. o, anterior; p, posterior; V, ventral; d, dorsal; be, blastocoele; gc, gastrocoele. (From ConJdin, "Heredity and Environment," Princeton University Press, A-H after Hatschek, by permission.)
FIG. 160.
I cell;

B, 2





form two

Shortly after fertilization has taken place, the zygote divides to cells that remain together (Fig. 160). Each of these

then undergoes a division in a plane at right angles to the


Then each cell divides in the one, and four cells are formed. third plane, resulting in the formation of dght cells. These early stages in the development of an individual are called cleavage


of cells



division continues until a small


mass cavity, which

and at the same time a central termed the cleavage
built up,



blastocoekj appears. now said to be in


blastula stage, a blastula being merely a hollow sphere consisting of a single

layer of cells surrounding a cavity.

The preceding account
cially to the

applies espeof eggs in


which the yolk is evenly distributed. In cases where a large amount of yolk is massed at one end of the egg, as in birds, reptiles, and most fishes, the cleavage divisions do riot extend the entire length of the embryo, but the portion with the yolk remains undivided (Fig. 161). In the arthropods, where the yolk is confined to the
(Fig. 170).

FIG. 161. Cleavage in the egg of the gar pike (Lepidosabout 5 hours after teus),

The yolk-laden






(From Skull,

Principles of Animal BiolOQV," after Eycleschymer.)

center of the egg, cleavage involves only the peripheral portion
Gastrulation. Following the formation of the blastula, a very important phase of development occurs (Fig. 160). The cells on one side of the embryo, which ordinarily are slightly larger than the others, begin to invaginate, or bulge inward, and this process continues until the lower and upper cells are in contact and the cleavage cavity is almost obliterated. The embryo is now called a gastrula. The new cavity formed by the process of
gastrulatien is known as the archenteron or gastrocoele, while the opening at one end of the embryo is the blastopore. The outer
layer of cells comprises the ectoderm, the inner layer the endoderm. As our study of the hydra has clearly shown, some- animals

go uo farther in their development than the gastrula stage. Both the sponges 1 and coelenterates have only two layers of cells
1 The outer and inner layers of a sponge seem not to correspond to the 3ctoderm and endoderm, respectively, of a coelenterate, since in the sponge's



surrounding a single cavity that communicates with the outOf course, many minor modificaside by means of one opening. tions in this fundamental plan arise, such as the development of tentacles in the hydra, but these are special features that are related to the life habits of the animal and enable it to carry on a




FIG. 162.


Cross sections of older embryos of Amphioxus in successive stages of development, showing the formation of coelom and mesoderm. ec, ectoderm; en, endoderm; mes, mesoderm; np, neural plate; nt, neural tube; nc, notochord; gc, gastroooele; cod, coelom; ent, enteron. (After Conklin, "Heredity and Environment," Princeton University Press, by permission.')



The most important




concerning the two

remain permanently In the higher animal groups, the gastrula now elongates somewhat, and further changes take place. The most important of these are concerned with the formation of the mesoderm and
development the position of the primary layers apparently becomes

lowest groups of metazoans is that they in the gastrula stage of development.

Mesoderm and Coelom Formation. In all metazoans except the sponges and coelenterates, there now arises between the two
primary layers of

may be derived from the ectoderm,

a third layer called the mesoderm, which endoderm, or both, depending

upon the l^nd of animal (Fig. 162). The ectoderm, endoderm, and mesoderm constitute the three primary germ layers. From them are later derived all the specialized tissues of the body. As previously pointed out, animals without a mesoderm (sponges and coelenterates) are said to be diploblastic, while those with a mesoderm (all other metazoans) are triploblastic. This is an
important distinction. Another noteworthy feature of embryonic development is the differentiation of the archenteron into a coelom and an enteron, the former developing outside the latter (Fig. 162). As previously pointed out, a coelom is characteristic of all metazoans
except the sponges, coelenterates, and flatworms.

Thus the

sponges and coelenterates have neither mesoderm nor coelom, the flatworms have mesoderm but no coelom, while the higher groups have both mesoderm and a coelom. The transitional
position of the flatworms, from this standpoint,

very apparent.

The methods among the different groups

mesoderm and coelom formation vary greatly

metazoans, but these details are

not important.

A common







Here the mesoderm

seen to arise from the endoderm

by the

formation of a pair of lateral pouches that become cut off, the archenteron thus giving rise to the coelom (surrounded by mesoderm) and the enteron (surrounded by endoderm). The


of the lancelet at this stage of

development also shows

the way in which the central nervous system of the chordates arises, viz., as a dorsal infolding of the ectoderm, the edges of

which unite to form a tube. The notochord is seen to arise as a dorsal outgrowth of the endoderm. This structure, it will be recalled, appears in the embryogeny of all chordates but is persistent throughout

Later Development.

only in the lower members of the phylum. The further growth of the embryo

complex and subject to great variation among the higher metazoan groups. Up to this point there is little or no differentiation between the cells of the embryo, no specialized tissues being formed as yet. In the subsequent development, however,
each of the three primary germ layers gives
rise to definite sets

of tissues.

In the vertebrates these

in part, as follows:

The ectoderm

gives rise to the outer part of the skin, to certain superficial appendages, such as scales, hair, feathers, nails, etc., and to the entire nervous system. The endoderm forms the
lining of the digestive

tracts and of t^e liver and gives rise to the muscles, connective and supporting tissues, blood vessels, the blood itself, and most of the other tissues of the body. Nearly all the organs of the

and respiratory


The mosoderm

adult animal are composed of
of the three
It is


derived from more than one

primary germ

layers of the


a remarkable fact that animals so diverse in their adult

stages as an earthworm, a starfish, a frog, and a mammal should begin their development in essentially the same way and follow a Minor variations are many, espesimilar sequence of stages.


but the main features of developonly in the later course of embryogeny in the formation of organs that a great deal of diversity arises.

beyond the

earliest stages,
It is

are constant.

Oviparity and Viviparity.
tion, the

In animals with external



necessarily develops outside the body.


fertilization is internal,

however, the embryo


develop within


the body, as in practically all mammals, or outside, as\'in all and most reptiles. In the two latter groups fertilization

occurs before the shell is formed, and so when the "egg" is laid, the embryo has already started to develop inside it. The large amount of food present is entirely consumed by the embryo during the course of its development. Animals in which the embryo develops outside the body are said to be oviparous, regardless of whether fertilization is external or internal. Except in a

very few





reptiles are oviparous, while birds are oviparous

amphibians, and without excep-

Nearly all the mammals, on the other hand, are vivipawhich means that the embryo develops within the body, deriving its nourishment by direct absorption from the maternal

Only two egg-laying mammals are known: the duckbill or platypus (Ornithorhynchus) and the spiny anteater (Echidna), both natives of Australia and adjacent islands (Fig. 163). The young are hatched from a large shelled egg like that of reptiles and birds. These forms, known as monotremes, are the most

Not only are they oviparous, primitive mammals in existence. but they have a cloaca, an organ characteristic of lower vertebrates but absent in all other mammals. Another primitive
feature of the



the fact that milk


secreted over

the entire ventral surface of the body, the mammary glands not being localized to form mammae (nipples) as in other mammals.

Intermediate between the monotremes and the higher


mals are the marsupials, a curious group including the kangaroos



Egg-laying mammals.

A, Duckbill (Ornithorhynchus) anteater (Echidna).


B, Spiny



many Australian relatives, and the opossums of America.

which have no shells, develop within the mother's is only a slight connection between the embryo but there body, and the wall of the uterus, no true placenta being formed. The young are born in a very immature condition and are immediately placed in an abdominal pouch (marsupium) located on the ventral side of the female. Within this pouch are the mammae to which the young become fastened. The following table is a convenient summary of the relations


discussed above.



Intrauterine Development. In all mammals except the monotremes and marsupials, the embryo, while within the body of the mother, receives its nourishment through an absorb-

FIG. 164. Diagrammatic section of human uterus with young embryo, a, dorsal wall of uterus; b, placenta; c, fluid-filled cavity surrounding embryo; d, lower ends of oviducts (Fallopian tubes); e, embryonic membranes; /, uterine tissue; 0, uterine cavity; h, embryo; t, umbilical cord with blood vessels. (From " The MacmUlan Company, by permission.) Woodruff, "Foundations of Biology

ing organ called the placenta, and thus nearly




The general features placental mammals (see pp. 187-189). of the female reproductive system of vertebrates have been illustrated by the frog, and it has been seen that the ripe eggs
escape from the pair of ovaries, enter the open ends of the

oviducts, and pass downward to the uteri where they are temporarily stored (see p. 177). In mammals the uterus is There may be either similarly an enlargement of the oviduct. two separate uteri (as in most rodents), the uteri may be fused

only at their lower ends (as in carnivores and many ungulates), or the fusion may be complete, resulting in a single undivided uterus (as in man and other primates). Fertilization occurs in one of the oviducts, the zygote passing down into the uterus where the embryo undergoes its development. At first the embryo uses up the food previously stored
in the egg,


but soon another source of nourishment becomes This is obtained from the mother by means of an organic attachment between the embryo and the uterus (Fig. A disk-shaped placenta, is formed consisting of both 164). maternal and embryonic tissues. It is firmly attached to the wall of the uterus and connected with the embryo by means of
the umbilical cord. Food and oxygen are brought to the placenta by the maternal blood vessels and pass to the embryonic tissues by diffusion. The blgod vessels of the embryo extend the length of the umbilical cord and into the placenta. They not only

absorb nourishment and oxygen from the maternal circulation, but give back carbon dioxide and other waste products.

The foregoing facts make it clear that the embryo is essentially a parasite in the body of the mother. It should be distinctly understood, however, that the blood vessels of the mother and the embryo are entirely separate from each other, and consequently no blood passes between them. All the blood cells
embryo, like all its other tissues, have been derived There is merely an osmotic movement directly from the zygote. through the placenta of food and oxygen from mother to embryo and of waste products in the opposite direction. There are also no nerve connections, and consequently there is no way in which the mental state of- the mother her thoughts, desires, or fears can influence the unborn offspring.
of the

Larval Stages. Until the individual becomes, independent of the mother or of the nourishment contained in the egg, it is an


it is born or hatched, as the case may be, it may developed and thus show a general resemblance to the adult parents, or it may be very immature and very unlike the parents. For example, when a grasshopper's egg hatches,


fairly well

bees. and usually consume Then they go large quantities of food during this period. grubs. an individual that bears practically no resemblance^ to the adult insect (Fig. These insects live for a relatively long time in the larval stage. butterflies and moths the young are called caterpillars.242 FUNDAMENTALS OF BIOLOGY is the young individual unmistakably a little grasshopper. Inc. 165). "Evolution and Life. and ants. egg.. New York. a. b." D. in the beetles. Appleton-Century Company. Finally they emerge .) 165. as full-fledged. Animal Metamorphosis of the monarch butterfly (Anosia plexippus). the young is called a other insects pass through a larval stage. In all Many cases similar to the latter. c. larva. pupa. wasps. larva. active adults. In the FIG. but a butterfly's egg hatches into a worm-like caterpillar. maggots. in which further development takes place. called the pupa stage. into a resting condition. (From Jordan and Kellogg. and in the flies. d. adult. by permission.

and reptiles) the eggs are laid without any further attention being given hatched they must great numbers of When the young are As might be expected. . formation of a larva into an adult is known as a metamorphosis. shift for themselves. Parental Care. an independent existence as only the amphibians that exhibit a A tadpole is really a larva.REPRODUCTION AND DEVELOPMENT IN ANIMALS 243 Among vertebrates it is larval stage in development. leading it gradually develops toward the adult condition. but remain aquatic throughout life. relatively few young are produced. but they are so protected that a much greater proportion survive. destroyed by other deleterious agents. 166. it is necessaiy that enormous numbers of eggs be produced. Some of the salamanders do not go beyond the The translarval stage. eggs and young are eaten by other animals or them by the parents. in the lower groups. In most of the birds and mammals. amphibians. on the other hand. In most invertebrates and in nearly all vertebrates belonging to the three lower groups (fishes. because few offspring reach maturity. Nest of ruffed grouse. Consequently. FIG.

In . degrees of lower groups. of course in the mammals the relation of the mother unborn young is much more intimate. snipe. quail. and mice are born blind and in a very helpless condition. prenatal development goes much farther and when hatched the young are provided with a covering of down. Thus in all our common songbirds the young are born blind. In all mammals the fact that the young are nourished by the mother's milk brings about a relation between parent and offspring that is absent in the The young of mammals similarly exhibit various dependence upon the parents. relying ground A cats. human young upon the long period of dependence of the the mother. merely depositing their eggs on the Others build a very crude nest that serves (Fig. They have to be raised in a nest of elaborate construcIn such birds as tion and be fed and protected by the parents. etc. few birds do not even incubate their to hold the eggs. cattle. and cannot live without parental care. without feathers.244 FUNDAMENTALS OF BIOLOGY In most birds the eggs are incubated by one or both of the parents. eggs. can walk. and unable to walk or fly. Some of these birds make no pretense at nest building. dogs. grouse. domestic fowls.. and guinea pigs are much more mature and are soon independent of parental care. and need little or no parental care. the heat of the body being essential to the growth of the embryo. and. both before and after birth. fact. to the birds gulls. For example. 166). is regarded as having been a very potent factor in the evolution of the higher mental faculties and of social organization. The young of most and mammals are very helpless at birth. while newborn horses. ducks. only for heat upon some external source. have eyes.

is progeny and the due to an actual organic continuity between successive generations. Fundamental similarity is the rule among organisms related by their parents. For the present we shall confine our attention to the great heredity reserving for subsequent chapters a consideration of the part that external factors play in determining the individual and racial constitution of organisms. but due to the direct transmission from one generation to the next of a material substance chromatin possessing the possidirecting the development of the individual along certain definite lines. both structural and from the interaction of two influences: heredity and environment. Certain like most of the resemblances. but usually most of them.CHAPTER XV PHYSICAL BASIS OF HEREDITY an organism. especially in their application to man. Every individual is unique. similarity may be seen. are due to heredity. All the characteristics of functional. arise are problems of great interest. In general. 7 but individual variability is apparent everywhere. Thus the characters that an organism bilities for internal influence comes to have are largely predetermined by reappearance of parental characters in the its ancestry. Although offspring tend to develop characters like those of a number of differences are always apparent. The fact that characters are transmitted through successive When a comparison is made between generations is self-evident. an individual and its parents. hereditary variations may represent new combinations of parental characters. descent. and the relative importance of each in the development of an individual. in a great many respects a marked This resemblance is not accidental. between parents and their offspring are due to environmental influences. Their relation to each other. while others may be due to the reappearance of Thus an individual ancestral characters latent in the parents. "like begets like/ but likeness is never complete. Some of the differences 245 . especially with respect to details.

an individual transmits all its hereditary characters to all its offspring. where inheritance is uniparental. Even though the egg ordinarily is immensely larger than the sperm. Here every individual is strictly biparental in origin.246 FUNDAMENTALS OF BIOLOGY differ from both of its parents in a given particular. It arises as a cell formed by the fusion of two gametes. Here the offspring is merely a detached portion of the parent. it will produce exactly the same kind of grapes as if it had been left to grow as part of the original vine. but. or by repeated self-fertilization. Uniparental Inheritance. differences exhibited It is the laws of heredity are essentially the same in all living thingvS. it is evident that an individual has only one parent. deals with the resemblances and by organisms that are related by descent. The biological science of genetics. Like other fundamental scientific generalizations. so but seeks heredity. For example. that occur of does related amount individuals variability among propagated asexually. and thus the complete The slight hereditary resemblance between them is explained. each coming from a different parent. if a cutting taken from a grape vine is planted. transmission. Where reproduction is asexual. due may to his inheritance. may be inherited. In such cases there is practically complete resemblance between the parent and its offspring. In all cases where sexual reproduc- tion involving two different individuals occurs. These facts make it clear that differences. both parents are equally potent in transmitting . a very different situation prevails from that which has just been described. but resemble one or more of its grandparents or even a more remote ancestor. and consequently the latter show not only almost perfect resemblance to the parent but also to one another. one of the most interesting fields of modern scientific study. possesses in a latent condition. composed of precisely the same kind of protoplasm. Biparental Inheritance. of all problems concerned with hereditary as far is possible. and so what is learned from a study of one kind of organism may be applied to others. as well as resemblances. or where it is sexual but self-fertilization always occurs. not concerned only with the known facts and principles of to arrive at an ultimate understanding. The foregoing facts make it clear that. arises for the most part under the influence of the environment. and that a parent may transmit characters that he himself does not manifest.

It is apparent. 2 The only exception occurs in the special case of identical twins. 2 if This is particularly true It is evident that. as will be seen later. biparenacteristics to any one of its offspring. All the organism's inherent capacities are present in the zygote. therefore. they are the conveyors of the heritage. from the standpoint of heredity. is explained (see also p. . capacities for developing along 1 The reason for this is that the physical basis of heredity is contained in the nucleus. but here the two individuals are produce^ by fission frt>m a sing1$ the same inheritance. formed through the act of fertilization. Bridge. zygote. present in the gametes that represents characters and makes It inherits potentialipossible their subsequent development. that a child cannot inherit any of its parents' actual peculiarities. that because the gametes are the sole means of maintaining organic continuity between successive generations. the parents belong to diverse racial stocks. It should be kept in mind that an organism's inheritance is To say that characteristics the sum of its innate capacities. having been contribution. Thus their resemblance. It is obvious or traits are inherited is to speak figuratively. some ties predetermined line. 247 In biparental inheritance an individual from one parent and some from the other. for each of the progeny may represent a different tal inheritance results in among combination of ancestral characters. it necessarily follows that this single cell carries the parent 's entire hereditary The Hereditary This means that the zygote. not as a unit. brought together by the fusion of the two gametes. an individual must be regarded. What it does inherit is something as these belong to the parents. and so have amounting to practical identity. inherits some of its peculiarities a much greater degree of variability offspring of the same parents than does uniparental inheritance. Since the only material contribution that each parent makes to its offspring is a gamete.PHYSICAL BASIS OF HEREDITY 1 hereditary characters. most of them independently heritable. must contain all the potentialities for the complete development of the new individual. which happened to have been brought together when the individual came into existence as a zygote. but neither parent transmits all its hereditary CharAs a consequence. and which will later become separated and redistributed in various ways to its own offspring. 260). but as an aggregation of innumerable characters.

metaphase. as follows: (1) A multiplication of cells occurs by cell division. In all multicellular organisms. and then certain matters closely associated with it. 167. growth presents three overlapping phases. A. (2) this is followed by a limited amount of cell enlargement. growth takes place cells tion of new from those already present. they derive from not only their living substance. In animals it is most active in embryos but continues in all parts of the body until the adult stage is . early telophase. //.000. B. resting cell. For this reason it is necessary to consider briefly. cell differentiation takes shall be place. a particular kind of tissue being formed. late prophase. (3) finally. by the formaIn general. F 6 H Successive stages in cell division in a root tip of onion (Allium cepa) X 1. which means their capacity to develop in a definite way. Cell division We in plants is confined largely to root tips. cell division completed. and to the cambium of roots and stems. F. The phenomena of hereditary transmission are closely associated with the process of cell division. first the mechanism of cell division. G. late telophase. but also their inheritance. When a cell gives rise to two new cells. buds. C. E. concerned here only with the first phase of growth. anaphase. it E FIG. D. early prophase.248 FUNDAMENTALS OF BIOLOGY Vegetative or Somatic Mitosis.

into a definite number of rod-like structures called chromosomes. some of difference which form the bipolar spindle. Another conspicuous between most plant and animal cells is that in the latter . Two groups of delicate fibers now appear in the cytoplasm at opposite poles of the nucleus. the nuclear membrane and the nucleolus gradually disappear. In animal cells. there are usually a pair of minute spherical bodies called centrosomes that are concerned with the formation of the spindle (Fig. up to the disappearance of the nuclear membrane. and the fibers meet to form a bipolar spindle. These constitute the polar caps. and that during which they move to the poles is the anaphase. From these bodies fiber-like radiations extend in all directions. chromosomes. while others extend from pole to pole. constitute the prophase. each group radiating inward When a cell is resolves itself. its own common center. 168). The halves of each split chromosome are now drawn to opposite poles of the spindle. a new cell wall forms on the spindle midway between to them. After the chromosomes have arrived at the poles. The initial stages of mitosis. and in some of the lower plants. cells divide by mitosis. cells. By this time each from chromosome has become longitudinally split into Some of the spindle fibers are attached to the two equal parts. the chromatin of its nucleus apparently by a process of condensation. each group of chromosomes gradually becomes organized to form a new nucleus. the telophase is reached.PHYSICAL BASIS OF HEREDITY reached. Figure 167. tip. apparently by a shortening of the spindle fibers that are attached reaching the poles. At the equator of the spindle the chromosomes become arranged in a plane perpendicular to its long axis. thus dividing the cell into two parts. preparing to divide. The stage during which the chromosomes lie at the equator of the spindle is called the metaphase. Upon the two daughter nuclei. in both plants and animals. cells. Meanwhile. At the same time. 249 Then it is limited principally to groups of unspecialized tissues. essentially similar showing a series of stages in vegetative mitosis as seen in a root should be carefully studied in connection with the following account. found in most which replenish wornout or injured Except in a few rare cases. a complex It is process in which the nucleus is conspicuously involved.

each chromosome splits longitudinally. 20 Lily. each of the daughter nuclei has a double set of chromosomes. in significance connection with hereditary behavior. 14 Onion. Mitosis represents an elaborate mechanism that secures an exactly equal distribution of chromatin to each daughter nucleus out and thus preserves a constant number of chromosomes throughThis is of the greatest all the body cells of the organism. B. 16 Indian corn. 60 Crayfish. 200 Significance of Fertilization. With FIG. anaphase. Then when the zygote undergoes its first division. X 1. as the following examples show: PLANTS Garden pea. the zygote receives two complete sets of chromosomes. 48 Horse. 48 Cotton. 24 ANIMALS Hydra. 18 Frog. 12 Domestic fowl.000. 144 Earthworm. 168. few exceptions. 32 Man. 56 Shield fern. as in an ordinary mitosis. As a result. Fertilized egg of a roundworm (Ascaris megalocephala) showing A metaphase. C. telophase. The centrosomes with their conspicuous radiations are characteristic of animal t cells. the number of chromosomes is definite and constant for each species of plant and animal.250 FUNDAMENTALS OF BIOLOGY the cytoplasm divides by a simple constriction rather than by the formation of a cell wall. 26 Tobacco. half of which are paternal . three stages in cell division. one from the sperm and the other from the egg. When two gametes unite. the halves passing to opposite poles of the spindle.

the other maternal. 169). the members of each pair are said to be homologous. cells. and eventually to all the somatic or vegetative cells of the adult Thus each body cell contains a descendant of every chromosome that was present in the zygote. one member of each For this pair being paternal in origin. 251 is Because this behavior repeated with each subsequent somatic cell division. half maternal (Fig. the other by the egg. B D FIG. and each pair may have its own structural individThe body uality. Reduction of Chromosomes. 258). the double chromosome number is transmitted to all the cells of the embryo organism. The reason. one set having been contributed by the sperm. size two homologous chromosomes forming each pair are alike in and shape (except in the case of the XY pair described on p. the gametes. It is important to understand how this reduction in chromosome number is brought about. Each of the two cells arising from the zygote has a double set of chromosomes. are designated as with a single set. as haploid. often differing conspicuously from other pairs. having a double set of chromosomes. 169. it contains a definite number of pairs of chromosomes.PHYSICAL BASIS OF HEREDITY in origin. E Diagram F of fertilization and cleavage. that is. diploid. It has already been stated that sperms and eggs in animals are derived from unspeciaiized cells .

FIG." chromosomes animals (Fig. the germ 1 involves two cell divisions. It is at the time of ovulation that the germ cells undergo maturation. ment proceeds. 170). and this ripening process. in most cases continues throughout the lifetime of the individual. 1 at the posterior end. that each unripe In the human male. dividing by the regular mitotic process to form a large number of spermatogonia in the male and oogonia of these in the female. the germ cells undergo a period of multiplication. normally takes place only between the ages of thirteen and forty-five. the latter having arisen from a single The yolk (y) primordial germ cell. the germ cells continue to multiply during childhood. During childhood the female germ cells increase in size and accumulate reserve food. a very early stage of embryonic development a primordial germ which all the germ cell from be derived (Fig. Some size. now increase in becoming primary spermatocytes and primary oocytes. When the animal has reached maturity. but in the female -the period of multiplication ends before birth. 229-231). The liberation of ripe eggs from the ovaries. and not from differentiated somatic tissues In some cases it is possible to identify in (see pp. but otherwise remain dormant. called ovulation. called maturation. in is the (From Shull. center. An early stage in the development of the embryo of the fly Miastor. much less numerous than the primary spermatocytes but be- come a great deal larger. showing the somatic cells (d) at the periphery and the germ cells (0c) some of the germ cells ripen.) " Reduction in the number of Animal Biology.252 called FUNDAMENTALS OF BIOLOGY germ cells. respecThe primary oocytes are tively. Principles of after Hegner. and it is supposed that one or two eggs are extruded every 28 days. cells will cells of the emthe other All bryo become specialized to form somatic tissues and take no part As developin reproduction. takes in directly 171). place in connection of with the formation cells of gametes Maturation is. . 170.

171. the primordial germ cells. come together in pairs cell SpcrmaJogenesis Oogenesis Sperma-j-^ '^TO //I y^ SpermaiocyfeE / \ 4- \ \ Mafurq-fion division -I x Cleavcfge FIG. all of which are diploid. This unique pairing of the chromosomes. the chromosomes. Like all the somatic the unripe germ cells are diploid. which takes place at no other time in the life history. At the time of the first cell division. and in cleavage. give rise to a large number of unripe germ cells (called spermatogonia in the male and odgonia in the female). and remain in contact until the bipolar spindle is formed. by an indefinite number of cell divisions (only two of which are shown in the diagram). Diagram showing the behavior of the chromosomes in the multiplication and maturation of the germ cells. in fertilization.PHYSICAL BASIS OF HEREDITY germ cells. after being formed from the chromatin of the resting nucleus. During the multiplication period. It is apparent that there are half as many chromosome pairs as there . 253 produces four potential gametes. is called synapsis.

poles of the spindle (Fig. Instead. 172). often before the two daughter nuclei have returned to the resting condition. Each chromosome of every pair now splits longitudinally. but unlike what takes place in an ordinary mitosis. the two halves of every split chromosome simply pass to opposite poles. have been derived. . but every chromosome is longitudinally split in preparation for the next mitosis.254 FUNDAMENTALS OF BIOLOGY were separate chromosomes in the unripe germ cell. The second division follows at once. there is no immediate separation of the halves. so that the four resulting nuclei have half as many chromosomes as were present It should be borne in mind that each in the unripe germ cell. When a spindle is organized in each nucleus. 172. each of the two daughter nuclei contains one member of every pair of chromosomes present in the nucleus from which they Fio. Diagram showing the difference between the behavior of the chromosomes during a somatic cell division (upper line) and a reduction division (lower line). the two split chromosomes constituting every pair merely separate and move to opposite Upon completion of the mitosis. of the four haploid cells that come from an unripe germ cell contains descendants of only one member of each pair of chromo- somes that were present in the unripe germ cell.

the larger one being the secondary oocyte and the smaller one the first oocyte always undergoes another unequal division. Thus each of the four cells derived from a primary spermatocyte becomes tids. with reference to its mate. the polar bodies soon disintegrating. are the same in both sexes. each pair of cells has a different assortment of chromosomes. since this depends upon the position in which each chromosome happens lie. in turn. promiscuous but always involves homologous chromosomes. Then. divides to form two spermaThese are transformed directly into sperms. in an animal where the diploid number of chromosomes is six. as outlined above. each primary spermatocyte gives rise to two secondary spermatocyteSj each of which. (1) At synapsis the pairing of chromosomes is not Each tion. Two very important details concerning the process of reduction remain to be considered. to four cells are produced by the regular mitotic division that always follows the reduction division. In this way functional. again. it is entirely a matter of chance to which pole either member of a pair passes. (2) When the separation of these homologous chromosomes takevS place during the reduction division. and other respects originally contributed by the egg. but the secondary The latter may or may not divide The egg alone is large egg and two or three small polar bodies. To illustrate. when they are lined up on the equator of the spindle preparatory to separation. during the previous act of fertiliza- and a descendant of a chromosome corresponding to it in size. Thus each primary oocyte gives rise to a single or mature egg. the paternal chromosomes may be designated as Ay B and C. Consequently each of the two resulting cells has a different assortment of paternal and maternal chromosomes. In the male. a slight difference occurs. the maternal ones as a. polar body (Fig. a functional sperm. All y when . form. of Grouping Chromosomes in Gametes. producing the second polar body and the odtid.PHYSICAL BASIS OF HEREDITY 255 Although the essential features of maturation. however. In the female. one cell among gets all the yolk that otherwise would be equally divided four cells. 6. the division of the primary oocyte results in two cells that are very unequal in size. 171). 173). and c (Fig. pair consists of a descendant of a chromosome originally contributed by the sperm. but only one member of each pair of homologous chromosomes can be present.

abC. The number of different kinds of gametes depends upon the number of chromosomes. Because the number of chromosomes is doubled in fertilization. B with 6. and abc.777. Assum- ing that the chromosomes are the bearers of hereditary elements. In other words. AbC. Number of possible kinds of gametes 4 In the case of man. Diagram chromosomes to the That the number of possible kinds of is gametes formed depends follows: on the number of chromosomes Pairs of shown as cells. A Be. the possible number of different kinds of gametes is 16. there are eight different ways in which the three kinds of chromosomes may be grouped: ABC. In synapsis. explain four things: (1) why tion . aBC. etc. a reduction must occur before or when gametes are formed in order that the number of chromosomes can remain constant through successive generations of diploid individuals. prevails among offspring of the same parents. where the number of chromosome pairs is 24. A pairs with a. Since the gametes can have only one member of each pair of homologous chromosomes.256 FUNDAMENTALS OF BIOLOGY the somatic cells and unripe germ cells contain three kinds of chromosomes and two chromosomes of each kind. and C with c.It is evident that this fact accounts for the diversity that 216. aBc. 2 chromosomes in unripe germ 3456 8 16 32 64. Gametes of parents Zygote and all aoma tic cells of offspring Kinds of gametes offspring produces illustrating the distribution of paternal and maternal gametes formed by the offspring. one paternal and the other maternal. 173. it follows that there can be eight possible kinds of gametes formed in approximately equal numbers. Abe. Flo. the association of paternal and maternal chromosomes in fertiliza- and their subsequent separation at the time of reduction according to the law of chance.

however. (2) why the male and female are equally potent in transmitting their hereditary characters. 174. 49-51). (4) why fertilization effects innumerable new combinations of ancestral characters. Lippincott Company. It introduces and diploid individuals. but in some of the thallophytes and in all the three . Alternation of generations in plants involves more than the production of two kinds of FIG. by permission. reduction of chromosomes always occurs directly in connection with the formation of gametes. the haploid number is carried over to all the cells of the gametophyte.) . (3) why the same combination of paternal and maternal chromosomes do not enter into the formation of every gamete that an individual produces. formed by the fusion of two gametes. plant bodies in the definite haploid life history (see pp. including the gametes. In animals. "Physical Basis of Heredity." J. Male and female fruit flies (Drosophtta melanogaster) (From Morgan. mitted to all the cells of the sporophyte. When spores are formed.PHYSICAL BASIS OF HEREDITY inheritance in 257 all organisms that reproduce by sex is biparental (except in cases of self-fertilization and parthenogenesis). B. so that each spore is haploid. is the first cell to have the This number is then transdiploid number of chromosomes. (he reduction of chromosomes takes place. The zygote. the gametophyte representing the former and the sporophyte the latter. Significance of Alternating Generations. When the spore germinates.

(After and There are two pairs of large curved chromosomes. the eggs all alike. 175. in the formation of of the straight one known as the . 174 MALE FEMALE X Morgan. Since all the body cells and unripe germ cells of the female have a pair of -X-chromosomes. of which the male has one and the female two. The sperms are of two kinds.} Y . while the straight ones. an insect that has been extensively studied by geneticists.and a Ycells of the fruit fly (Drosophila melanogaster) chromosome correspond to the pair of X-chromosomes of the female. The sex of any one of the offspring depends upon the kind of sperm that entered into the formation FIG. This is F-chromosome. of the zygote from which it has developed. the end Puenta Gametes of parents Offspring Diagram showing the mechanism of sex determination in the fruit fly. 175). Diagram showing the chromosomes that occur in all the somatio In the male an X. Determination of Sex. the spores being formed in groups of four. it follows that. XX Fio. In the fruit fly (Drosophila melanogaster). 176. are called X-chromosomes. one pair of very small ones.258 FUNDAMENTALS OF BIOLOGY it higher plant groups takes place when spores are produced. chromosomes is slightly hooked. A slight visible difference exists between the sexes in that. and one pair of straight chromosomes about two-thirds as long as the curved ones. each of the body cells has eight chromosomes (Figs. in the male.

and for others. The on the other will form two kinds of sperms: half of male. the latter into a male. and thus the number of male female offspring will tend to be equally numerous. that future research will in plants show that the causes underlying sex determination and animals are practically the same. For example. but in the male the F-chro- very small. It now seems certain. Occasionally the F-chror mosome is absent. the female has 22 chromosomes and the male only 21. The former develops into a female. or there may not be any apparent difference at all. it seems probable that it is not the presence of the F-chromosome in the zygote that causes it to develop into a male individual but the presence of only a single ^-chromosome. if fertilized by the other kind of sperm. however. 176). many XX- produce as and There are a great many other kinds of animals in which a F-chromosome has been identified in the male. the missing one being the F-chromosome. the A -chromosome in the body cells of the male then being unpaired. each egg will contain 259 a single Ji-chromosome. So here sex is determined by the chromosome equipment of the egg. Because the two kinds of sperms are equally numerous. those of the male being all alike. the result is an XX-zygote.PHYSICAL BASIS OF HEREDITY of gametes. in birds and in moths it has been slightly different. in a few animals the situation is For example. There is reason to believe. In only a few plants where male and female individuals are differentiated has a visible chromosome difference been is mosome discovered. Consequently all the eggs and half of the sperms have 11 (10 plus an -chromosome). that both sexes have 48 as the diploid number. If an egg is fertilized by a sperm of the first type. in the squash bug (Anasa tristis). the rest of the sperms having only 10. In some cases it differs from its mate in size rather than shape. but the mechanism is exactly the same as where the male produces two kinds of gametes. Although most cases of sex determination follow the same scheme as that described above. For this reason. . random mating of gametes will as XF-zygotes. them will have an X-chromosome and half a F-chromosome. hand. as a result of very careful recent investigations. however. found that the female produces two kinds of gametes. an XYzygote is formed (Fig. X In the case of man there has been a diversity of opinion in regard to the total number of chromosomes and to the difference between the male and female sets.

regard to they . and those which arise from the splitting of an embryo in an early stage of cleavage (identical twins). This is because have exactly the same chromosome equipment. however. are invariably of the same sex and are alike in always of the same sex.260 FUNDAMENTALS OF BIOLOGY of That the sex an individual is determined at the time of fact that identical twins are fertilization is substantiated by the Human twins are of two sorts: those coming from two different zygotes (fraternal twins). and are no more alike in their hereditary characters than ordinary brothers and sisters born at different times. The latter. The former may or may not be of the same sex. both individuals having arisen from the same zygote. all their other hereditary characters.

crossed certain varieties of garden peas and discovered that He their differentiating characters are inherited in accordance with Mendel published his results in definite mathematical laws. FIG.CHAPTER XVI MENDELIAN LAWS OF HEREDITY laid The foundation for our present knowledge of heredity was by the work of Gregor Mendel. an Austrian monk (Fig. 177. 1822-1884. Biologists have since found that the laws of heredity discovered by Mendel apply to all organisms. when it was brought to light and its great -iu!ii(i'-:i:v appreciated. Gregor Mendel.i 261 . the unripe stamens it to the pistils of the other. . but unfortunately his paper attracted little attention and remained unnoticed until 1900. 1866. including man. and thus are of fundamental importance. 177). Two plants are crossed by taking pollen from one and applying In most cases.

They may distinctly different types of indito the same species or to different belong but ordinarily individuals that are not rather closely As an related cannot be crossed. is crossed with a white-flowered plant. when of the second flower must first cannot occur. for the sake of simplicity. that is. 178. piagram of a cross between a red-flowered and a white-flowered four o'clock (MirabUis). showing the history of the chromosomes carrying the genes for red (#) and for white (r) flower color. In either case. A hybrid is simply an organism whose parents represent two viduals. no offspring will result. the resulting seeds . 178). The hybrids are intermediate between the parents. red (pollen being taken from the former instead of the latter). illustration of Mendel's first law. When A the seeds are planted they give rise to hybrids with pink flowers If the reciprocal cross is made. however. planted. When these hybrids are allowed to pollinate themselves. species. give rise to hybrids. the hybrid plants are intermediate between the parents. shall choose a different kind of plant from the one with which he experimented.262 FUNDAMENTALS OF BIOLOGY be removed so that self-pollination The seeds produced as a result of a cross. the result is exactly the same. having red flowers four o'clock (Mirabilis jalapa) Principle of Segregation. that is. we Parents Garnet* of parents Offspring FIQ. white with (Fig.

and speckled (called "blue"). and 25 per cent white (Fig. and 50 per cent of their offspring are uals. of interbreeding pink-flowered four o'clocks. It is therefore apparent that pink-flowered four o'clocks are always hybrids. 179). Andalusian fowls: black." which means that when self-pollinated they produce only Similarly the white-flowered plants give plants with red flowers. The red-flowered plants derived from the pink parents " breed true. but when blue fowia A among animals of There are three kinds .MENDELIAN LAWS OF HEREDITY 283 give rise to three kinds of individuals in the ratio of 1:2:1. FIG. In other words. fact that each of the hybrid parents produces Diagram illustrating the results The 1:2:1 ratio arises from the two kinds of gametes. 179. always produce offspring in the ratio of 1 red: 2 pink:l white. The pink-flowered individ- Fferaote Grata of Offspring Ratio The principle of segregation. or vice versa. airways give rise to blue offspring. approximately 25 per cent of the plants will have red flowers. likewise hybrids. 50 per cent pink. similar case is seen in Andalusian poultry.Blacks crossed with whites. however. white. rise to only white-flowered progeny.

Hereditary Elements. as in the red (RR) and white (rr) four o'clocks. each carrying an R gene. The explanation of Mendel's principle of segregation is seen in the behavior of the chromosomes during fertilization and reduction (Fig.264 FUNDAMENTALS OF BIOLOGY is 1 are interbred. half only the r. Similarly all the gametes of the white four o'clock have a gene for white flower color (or lack the red gene. As a result. Hereditary characters are represented in all the cells of an organism by invisible units or elements called genes. the result black: 2 blue : 1 white. All the gametes produced chromosomes a gene This may be designated as R. the members of the pair of alternative genes must also separate and go to different cells. But if the two genes in each body cell are different. In other words. the haploid number. since a gamete can have only on3 member of each pair of homologous chromosomes. amounts to the same the formula rr. Blue Andalusians are hybrids exactly comparable in their behavior to pink-flowered four o'clocks. it can carry a gene for only one member of each pair of genes. and so the plant is designated by a red four o'clock is crossed with a white one. the zygote from one gamete a chromosome bearing an R gene. if they are contrasted or alternative in their relation to each other. the individual is said to be homozygous for the character in question. Because homologous chromosomes always separate when the reduction division occurs. from the other a homologous chromosome having an r gene Thus the pink hybrid is designated Rr because all (Fig. will have red it the same. as in the pink (Rr) four o'clock. 178). if both members of a pair of genes are the same. . its vegetative cells contain one gene for red (R) and one for receives When white (r) flower color. which are associated with the chromosomes. thing). In the four o'clock the gametes have eight chromosomes. When an egg and a sperm unite. the individual is termed heterozygous. designated RR. 178). In regard to any given character. which by a red four o'clock carry on one of their for red flower color. on the average. the zygote receives two R genes and all the vegetative cells derived from Hence the plant. that is. flowers. there are eggs and sperms) formed two different kinds of gametes (both by the hybrid in approximately equal numbers: half of them have only the R gene.

50 per cent pink (Rr) > 50 per cent pink (Rr). or when self-pollination occurs. as has been stated. Gene Combinations. white. These are called backer osses.. out of 100 trials there would result approximately 25HH. number of With trials. and thus two kinds the 1:2:1 ratio is explained.kinds. those involving a pure (homozygous) and a hybrid (heterozygous) individual.. and 25hh. as head (H) and tail (A). 50 per cent white (rr) All the preceding crosses have been considered except the last two. when brought together by two : fertilization. but those of the hybrid are of two different '. combinations (R with R. Thus there are only two kinds of . coexist in all the cells of the and later become separated in their own gametes without having had any effect on each other. 50 X X red (RR) white (rr) per cent pink (Rr). r and r with R) give rise to the same kind of zygote (Rr). The situation just described comparable to the simultaneous Referring to the two sides of the coin tossing of two coins. there is a random mating of two kinds of sperms (R and r) with sible This results in four posof eggs (R and r) (Fig. as follows Red (RR) White (rr) Red (RR) Pink (Rr) Pink (Rr) Pink (Rr) X red (RR) . six matings are possible. 179). which occur in the same frequency. the gametes pair according to the law of chance. It is is more closely approximated the and pink four : Backcrosses. one is likely to occur as often Of these four possible gametic unions. The gametes of the pure individual are all alike. In other words. 5QHh. because these combinations are governed entirely evident that the 1:2:1 ratio greater the different by the law of probability. o'clocks.MENDELIAN LAWS OF HEREDITY The essential feature 265 may be stated as follows a pair of gametes uniting in offspring of MendePs principle of segregation The genes representing each member of contrasted characters. R with r. and r with r). When two pink plants are crossed. 25 per cent white (rr) -> 50 per cent red (RR). Pink-flowered four o'clocks produce two kinds of sperms and two kinds of eggs. 179 should make these points clear. A is careful study of Fig. two (R with as anyother. red. namely.100 per cent red (RR) X white (rr) 100 per cent white (rr) X white (rr) -> 100 per cent pink (Rr) X pink (Rr) -> 25 per cent red (RR). r with R.

180. among the resulting . thus resulting in a 1 1 ratio among the progeny. peas are crossed with dwarf peas.266 FUNDAMENTALS OF BIOLOGY zygotes that can possibly be formed. or vice versa. In other words. not intermediate. Diagram illustrating the results of crossing a pink-flowered with a white-flowered four o'clock. That half of the offspring resemble one parent and half the other can be easily understood by reference to Fig. interbred (or allowed tallness Now when the dominant character the tall hybrids are to self-pollinate). one gene for tallness produces the same effect as if two genes were hybrids are tall Mendel designated present. : The all tall. 181). Only one of the parents produces two kinds of gametes. consider one of Mendel's If tall Dominant and Recessive Characters. Although genes for both characters are present in all the plants of the hybrid generation. and dwarfness the recessive. and these must occur in the same frequency. 180. but no shorter than the parent (Fig. FIG. We are now ready to own experiments with garden peas. the resulting Gametes <rf parents Offspring Ratio backcross. one manifests itself to the total suppressi9n of the other.

Fio. Dominant and . Diagram of a cross between a tall and a dwarf pea. a 3: 1 ratio results when they are interbred. showing the history of the chromosomes carrying the genes for tall ness (T) and for dwarf ness (t). Because the hybrids are similar to the dominant parent. 181.MEN DELIAN LAWS OF HEREDITY 267 Cfcnetesef parents Hybrids Gametes of hybrids Offspring of hybrids Ratio recessive characters.

The principle of segregation. will exhibit the it If homozygous. tell whether an organism is homozygous or heterozygous for a Gametes of parents Offspring Ratio FIG. but. this 3 :1 ratio is really only a modification of the 1:2:1 ratio. only them will (Fig. heterozygous.268 FUNDAMENTALS OF BIOLOGY progeny there are approximately three tall plants. howThe following list ever. if heterozygous. 181 clearly shows. smooth seeds over wrinkled. 182). however. is fundamental and always holds true. and hybrid (heterozygous) tall (Tt). Although the pure and hybrid tall plants are similar to each other on the basis of outward appearance. and found that purple flowers are dominant over white. Mendel experimented with other characters in peas. The backcross where dominance is showing the dominant character recessives. The phenomenon of dominance applies to the characters studied by Mendel and to many characters in other organisms but does not apply to most hereditary characters. as Fig. 182. to every dwarf. yellow seeds over green. to an individual showing the all the offspring half of dominant character. contains a few characters exhibiting dominance chosen from many that are known. while the dwarfs (tf) necessarily are always pure. It also shows how widely applicable are the laws of heredity discovered by Mendel: . It is apparent that there are two kinds of tall individuals pure : (homozygous) tall (TT). Because the parent occurs. half of the offspring are given character by backcrossing contrasted recessive character. Because the hybrids are visibly indistinguishable from the dominant parent. they do not have the same breeding Where dominance occurs. it is always possible to possibilities. etc.

. When be discovered a second important principle. 184). thus far have been considered are comparatively simple.MENDELIAN LAWS OF HEREDITY 269 The cases of inheritance that Principle of Free Assortment. for in regard to just one pair of concrossed. they give rise to monohybrids. The fact that two of these combinations are new shows that the two pairs of characters are inherited independently of each other. If a pea having yellow smooth seeds (YYSS) is crossed with one having green wrinkled seeds (yyss). four (Fig. different kinds of individuals appear in approximately the following proportions: 9 yellow smooth: 3 yellow wrinkled: 3 green smooth: 1 green wrinkled (Fig. all the hybrids will have yellow smooth seeds (YySs) because both of these characters are dominant When these hybrids are interbred. and as a result all of them the parents differ in trasted characters. Mendel also crossed peas differing with respect to two pairs of contrasted characters. thus obtaining dihybrids. however. 183).

Therefore the ntimber of peas that are both yellow and smooth is determined by multiplying the number of yellow peas by the number of smooth ones (3X3). referring to Fig. The hybrid produces four different kinds of gametes. 183. at the time of the reduction division. there must also be free assortment of genes carried . while the geries S (for smooth seeds) and s wrinkled seeds) are associated with another pair. 183.270 FUNDAMENTALS OF &IOLOGY In order to understand how this ratio arises. It is evident that the ratio of yellow peas to green ones must be 3:1 and of smooth characters peas to wrinkled ones the same. yellow smooth and a green wrinkled pea. Since. of green wrinkled 1 X 1. Similarly the number and of yellow wrinkled is 3 X 1. Diagram of a cross between a Principle of free assortment. of green smooth 3X1. The two pairs of contrasted characters are represented by genes borne on two pairs of homologous chromosomes. Parents Gametes of parents Offspring Gametes of v^x' offspring FIG. each pair of must be considered separately. it can be seen that the genes Y yellow seeds) and y (for green seeds) are associated with one pair of homologous chromosomes in all the vegetative By (for cells of the hybrids. paternal and maternal (for chromosomes are distributed to the gametes independently of one another.

which. Although each class includes individuals that look alike. random mating of gametes results in 16 possible unions. while those which look alike. 272. two kinds of yellow wrinkled. Ys. giving rise to the phenotype ratio of 9:3:3:1. belong to the same phenotype. regardless of whether they have the same hereditary constitution. but just one green wrinkled. Individuals with the same hereditary constitution are said to belong same genotype. Consequently the dihybrid peas produce four different kinds of gametes in approximately equal numbers. fall into four classes (Fig. and ys. Thus there are four Sperms SMOOTH YELLOW SMOOTH YELLOW SMOOTH YELLOW SMOOTH O O O O O O O YYSS 'YySS YySs YELLOW SMOOTH YELLOW SMOOTH YELLOW SMOOTH YELLOW WRINKLED YELLOW SMOOTH YELLOW WRINKLED YYSs YYss YySs Yyss YELLOW SMOOTH GREEN SMOOTH YySS . These are given on p. but only four phenotypes. 184). 184. all do not have the same breeding possibilities. Thus from the dihybrid cross there are nine distinct genotypes. of green smooth. . two kinds to the kinds of yellow smooth individuals. on the basis of outward appearance. resulting The dpiybrid ratio Diagram showing the 16 gametic unions from random mating of four kinds of sperms with four kinds of eggs. When two dihybrids are crossed. yS. It should be noted that only 4 individuals out of the 16 are 'homozygous for both characters. as follows: YS. YySs yyss YELLOW WRINKLED GREEN SMOOTH GREEN WRINKLED YySs Yyss 'yySs FIG. and each carries a different assortment of genes.MENDELIAN LAWS on different chromosome OF HEREDITY 271 pairs. and consequently these alone will breed true.

the offspring. When these are interbred. all showing dominance. approximately in the ratio of 9 rough black: 3 smooth black: 3 rough albino:! A smooth albino (Fig. rough or resetted coat is dominant to the ordinary smooth coat. as each parent contributes one dominant and one recessive character. are precisely the same in Fig. are white fur.272 FUNDAMENTALS OF BIOLOGY may be illustrated among from the chosen animals by an example heredity of guinea pigs. such as in regard to sthoothness of coat. White animals colored or either is mated with an stock from a derived black If a pure guinea pig principle of free assortment albino. but if the parents differ in another way. Thus \yhen a smooth black guinea pig (rrBB) is crossed with a rough albino (RRbb). When two trihybrids are . exhibit the dihybrid ratio. all the offspring will ^e black. may have many called albinos. like other domesticated These animals. and their genes are borne on three different sets of relations of these t^ one another as those of the four classes of peas shown The chromosomes. When there are three different pairs of contrasted characters. the hybrid offspring will be rough black (RrBb). 184. the resulting trihybrid produces eight different kinds of gametes. as colored fur is dominant If the parents are alike in other respects. forms. when interbred. 185). the simple to white. four types appear among the progeny. The monohybrid ratio of 3:1 arises when the hybrids are interbred.

there are 64 different gametic unions possible. 185. after Castle.) plexity of the hereditary mechanism.MENDELIAN LAWS : : OF HEREDITY 273 crossed. produce offspring in the ratio of 9 rough black: 3 smooth black: 3 rough white: 1 smooth white. (From " Baur. Einftihrung in die experimentdle Vererbungslehre. . the combinations is 1. I Results of crossing a smooth black with a rough white guinea pig. when it is realized that most individuals are heterozygous for many characters. we can appreciate why there is so much diversity among individuals of the same species." GebrUder Borntraeger. by permission. even among offspring derived from the same parents.048. giving rise to the phenotype ratio of 27 9 9 9 3 3 3 1 In the case of : : : : : . Furthermore.576. when interbred. Berlin. The hybrids (Fi) are rough black and. 10 pairs of differentiating characters. This gives number of possible some idea of the com- 3 FIQ.

All yellow peas would be smooth and all green ones wrinkled. "Principles of Animal Biology" after Morgan. normal red eye. G. bar eye. Then. Z>. (From Shull. vestigial wing. however. when the two chromosomes separate. beaded wing. in if the genes Y and S were borne on one chromosome. K. each has received from the other a group of genes that it did not have before. Free assortment occurs where different sets of genes are associated with different chromosome pairs. club wing.) or green smooth. A. F. and y and s on another. linked. be no new combinations such as yellow wrinkled FIG. there is no free peas. However. that is. ei al. Linked characters tend to be inherited together because their genes are carried by the same pair of chromosomes. 175 and 186). B. eyeless. borne on the same chromosome.274 FUNDAMENTALS OF BIOLOGY Linkage. Although the genes in different . /. 187). indicating that each chromosome bears not one but many genes. H. 186. rudimentary wing. E. all of the genes which have been studied fall into four groups corresponding to the four pairs of chromosomes (Fig. Some of the heritable variations that have arisen in the fruit fly (DrosophUa mdanogaster). This condition is called linkage. in pairs. notch wing. C. miniature wing. in the fruit fly. under ordinary conditions there could of characters. When homologous chromosomes are associated reduction division. There is a great deal of evidence." linked genes often become separated from one another and come to lie on different chromosomes. owing to an occasional peculiarity in the behavior of the chromosomes at the time of the crossing over. and yet the animal has only four pairs Where two genes are of chromosomes (Figs. /. It has been found that. For example. Thus. in the fruit fly about 400 different char- acters have been studied. white eye. Z/. normal wing. called " sometimes a portion of one is interchanged with the corresponding portion of the other. assortment. and so the inonohybrid ratio results. truncate wing.

0 085 1010 t050 FIG. PURPLE.0 SPINELESS - 645 59. 54.WHIT E-OCEltl HUMPY ARC "0PLEXUS BROWN SPECK 8o5. Moreover. 253 SEPIA = 2ao- : CUT 25. to determine the relative position of the genes on each chromosome with reference to one another. by permission.) of Actually. 187.. I. the numbers represent the distance of the locus of each gene from one end of the chromosome.VERMILION 33.< OARNETJ aO- 42JO - SCARLET PINK 'BLACK 455- ^2^ FORKED 570" ^BAR 465. The names II. through hereditary behavior. "Evolution and Genetics. showing the location seme of the genes that have been studied.0 - BITHORAX GLASS 650 - CLEFT 6SO ?ao 615 VESTIGIAL 656 675 LOBE - -DELTA.-ROUGH 057 101. III. -HAIRLESS r ' 680 --BOBBED. there is considerable evidence indicating that the genes on each chromosome are arranged in a linear IL 00 -ECHINUS 75. those in the same group show linkage.7 HI STAR QO + ROUCHOID TRUNCATE CROSSVT<L*S MJO J- i STREAK. corresponding to the four pairs of chromosomes.-CURVED 720. (From Morgan. ." Princeton University Press. Chromosome map of DrosophUa mdanogaster. and IV. as series. in the fruit fly it has been possible.MEN DELIAN LAWS OF HEREDITY 275 groups show independent assortment.1 MF** 2fto DACHS SKI 4.S13. These fall into four groups.-RUBY 5.8 + 3310 36. indicate the characters in the fly with which these genes are associated. EBONY 735.0 ^MINIATURE 315 -DICHACTE r I- SABLE : X 4.

has been one modern genetics.) Sex-linked Inheritance. 187. In many animals there are hereditary characters that are transmitted in a unique way. This is due to the fact that their genes are carried by the ^-chromosomes.276 FUNDAMENTALS OF BIOLOGY Proof that each gene is always situated indicated in Fig. 188. of the outstanding accomplishments of Parents Gametes of parents Hybrids Gametes of hybrids Offspring of hybrids FIG. on a particular chromosome. Diagram of a cross between a white-eyed male and a red-eyed female fruit fly. illustrating sex-linked inheritance. (Adapted from Morgan. . and at a definite locus on the chro- mosome. The genes for eye color are associated with the X-chromosomes.

but invariably the . 188). 189. 277 The inheritance of white eye color in the fruit fly is a well-known example. red-eyed male crossed with a white-eyed female fruit fly. (Adapted from Morgan. showing that red is the Gametes f Hybrids Gametes of hybrids Offspring of hytoi* FIG. all the offspring are red eyed. When a male fruit fly with white eyes is mated with a red-eyed female. 188. while half of the male are red eyed and half white eyed (Fig.MENDELIAN LAWS OF HEREDITY Such characters are termed sex linked. The typical monohybrid ratio of 3:1 appears. all ijbe female progeny have red eyes. however.) A dominant character. the reciprocal of the cross shown in Fig. When these individuals are interbred.

When the reciprocal cross is made. Diagram of the inheritance of human color blindness through the male. Diagram of the inheritance of human A color-blind woman has color-blind sons female. : : : : The type of sex linkage described above occurs in many other insects. and that the F-chromosome does not carry any genes. all the sons have white eyes Parents Gamete* of parents Offspring FIG. but and all the daughters red eyes (Fig. 190. 189). the latter are heterozygous for the defect. But when these hybrids are interbred. cats . a red-eyed male mated with a white-eyed female. that is. Gametes of parents Offspring FIG. in sheep. These results are easily understood if it is assumed that the gene for red eye color is carried by the X-chromosomes.278 FUNDAMENTALS OF BIOLOGY white-eyed individuals are males. The offspring are all phenotypically normal. their offspring fall into the following classes 1 red-eyed female 1 white-eyed female 1 red-eyed male 1 white-eyed male. colr blindness through the and normal daughters. but the daughters are heterozygous for the defect. in certain fishes and amphibians. 191.

having but one gene for the defect." A color-blind woman. mated the at to a normal man. but are merely heterozygous. . (the males are either by F-chromosome being not involved normal or actually color blind. Because the gene for color blindness X-chromosome all). fly. females may for the character. Since the character is recesvsive. mated to a normal woman. A color-blind man. because it depends upon the mating of a color-blind man to a woman who is either herself color blind (homozygous) or a carrier for the defect (heterozygous). 191). X Color blindness in man is a sex-linked character that follows the same scheme of inheritance as white eye color in the fruit Color blindness is recessive to its contrasted character. and probably in most mammals. in birds. the female only one (see p. the inheritance of sex-linked characters is exactly the reverse. or heterozygous Color blindness in women occurs in aoout 4 cases per 1. while carried be either normal. normal none of the daughters arc themselves color blind. and in some fishes. In butterflies and moths. because here the male has two -chromosomes. vision. and this seldom happens. 190). Such women are called "carriers. color blind. but to none of his sons (Fig. transmits the defect to all is her sons (Fig. 259).MENDELIAN LAWS OF HEREDITY 279 man.000 for men. transmits a gene for the defect to all his daughters.000 as compared with 40 cases per 1. on the other hand. Color blindness in women is very rare.

as has been done. Consequently an elementary course it is not possible to present more than a very general background of information pertaining to heredity. The mechanism of in transmission has been studied hereditary many different organhas been found and it that the fundamental features of isms. is in PLANT AND ANIMAL BREEDING The improvement of plants and animals must have started soon after primitive man began to raise crops and to bring wild animals under domestication. it has been on a scientific basis only since our modern knowledge of genetics has developed. Common observation demonstrates that of organisms all kinds in show considerable diversity 280 among themselves. Plant and animal breeding is an extensive subject that can be discussed here only in its barest outline. For example. Mendelism are practically universal in their application. But although the art of breeding has been practiced from time immemorial. and as a result a large mass of facts has been accumulating ever since. . however. Another complication arises from the fact that the mode of expression of characters often subject to environmental influences. Variation. a great impetus was given to the study of heredity. or to touch upon its application to human interests except in a very superficial way. while in other cases one gene may control the development of several characters. that heredity is a much more complex matter than our elementary consideration of it has indicated. It should be understood. many characters are determined by several or many genes which interact with one another to produce a given result. giving attention chiefly to the methods that modern scientific breeders use in developing improved races.CHAPTER XVII APPLICATION OF HEREDITARY PRINCIPLES With the rediscovery of Mendel's laws in 1900.

color of flowers. brighter flowers. selection process is continued with improved race of plants is developed. there is practically no evidence that they are transmitted to subsequent generations (see pp. by reason of this fact of universal variability that it is possible for plants and animals to be improved under man's guidance. and. Mass culture involves breeding from a plants. yield. Although it may seem that all variations are capable of being transmitted from parent to offspring. the following may be mentioned: vigor and rate of growth. Mass Culture. as well as a number of other crop plants. chief methods of selection human needs have been developed by or a combination of the two. Although such induced changes may be profound. size and shape of parts. it method of plant breeding. in The two common use are mass culture and pedigree culture. 281 no two individuals It is of any species are exactly alike in all respects.APPLICATION OF HEREDITARY PRINCIPLES fact. as they alone make racial improvement possible. larger fruits. The differences between individuals are called variations. its limitations should be kept in mind. This is a method of selection used It is the oldest only wiUi although has been more or less replaced by other methods. quality of fruits. 389- On the other hand. culture has certain advantages. etc. variations that are inherent that 393). disease resistance. To the latter class belong most differences that arise through the direct action of environmental influences and through use and disuse. such is not the case. Seed is collected from these superior individuals and sowed en masse. that are important to the breeder. The multitudinous races of plants and animals that serve selection. Of the great host of characters in cul- are tivated plants and domesticated animals that are determined largely by heredity. are determined by the germinal constitution of the organism It is only heritable variations of course transmissible. milk production in cattle. and egg production in poultry. The each generation until an Mass culture has been widely used in the breeding of corn and Although mass cotton. such as greater yield. hardiness. selected group of individuals. a group that varies in some desirable direction. it is still of considerable value. These are as follows: . drought resistance. hybridization. and consequently the existence of both heritable and non-heritable variations is recognized.

is often deceptive. permanent racial value. FUNDAMENTALS OF BIOLOGY In most cases it is necessary to continue the process of selection indefinitely. This method of breeding has long been applied to animals. owe their superior and such variations have no Mass culture is a slow method. but only rather recently to plants. 2. in many cases the offspring of the originally selected individual show so little All that is variation that further selection is unnecessary. cases. the others being discarded. each of which is made the basis of a separate race. 3. Mass culture cannot produce new characters but merely improves such characters as already exist. 6. Selection is made entirely on the basis of outward appearance.282 1. if isolated. or pedigree. double flowers. in many qualities to environmental factors. in heterozygous individuals. required is to prevent the plants of the improved race from crossing with inferior stock. An exact record. 4. The selected individuals. otherthe progeny will be superior. The pedigree method of breeding results in a much greater degree of uniformity among the members of the improved race than where mass culture has been used. is kept of the progeny of each selected individual. It involves the selection of single plants. and after several generations the best strain is preserved and made the basis of an improved variety. Thus selection is made on the basis of hereditary behavior. otherwise the improved strain deteriorates. Examples of mutants are plants bearing seedless fruits (which must be propagated vegetatively). It does not isolate the best individuals but merely raises the average quality of a large group. Pedigree Culture. In fact. as has been seen. not merely on outward appearance. The new variety will breed true if the plant that was originally isolated is homozygous for the superior it is desired to preserve. the progress made in each generation being slight. . 5. Seeds from each selected individual are planted in an isolated plot in order to prevent intercrossing. which.' characteristic or quality that all wise not 7 different from the others individuals that appear spontane- In nearly all cases they breed true for their peculiarities ously. Pedigree culture is valuable as a means of preserving mutants " This term is applied to individuals strikingly or sports.

for . with loss of pigment (albinos). the isolation of individuals and the development of homozygous strains result in a marked loss of vigor.APPLICATION OF HEREDITARY PRINCIPLES 283 purple leaves. produced by mating a jackass with a mare. The advantage is obvious. as a rule. In some cases. as then the hybrid can be propagated without using seeds." Pedigree culture is best adapted to plants that are normally self-pollinated and has been used very successfully with such With plants as wheat. etc. much inbreeding is Consequently necessary as a means of preserving a particular set of characters. oats. corn and certain other cross-pollinated plants. Although. only closely related organisms can be crossed. all members of an improved stock are usually more or less closely related. defined as an organism whose parents represent two distinctly different types of individuals. the best-known example being the mule. It is apparent that in animal breeding two individuals must necessarily be selected as the basis for an improved strain. be an index of its breeding possibilities. the progeny from a wide cross are sterile. The chief value of hybridization is that it brings together characters to form combinations that did not occur previously. however. usually a great many crosses of the same kind have to be made. peas. beans. Knowledge of the hereditary constitution of the selected individuals is available in their recorded pedigree. The method of hybridization is used extenA hybrid has been sively in both plant and animal breeding. etc. without horns. resulting Because a desirable combination depends upon chance. and animals without tails.. the newer method is Its appearance may or may not to consider its relatives as well. Although it is generally not advisable to mate the offspring of a single pair of parents. Many new varieties of plants and animals have arisen as "sports. Hybridization is particularly useful as a method of plant breeding where vegetative multiplication is possible. many hybrids have been produced by crossing individuals belonging to distinct species or even to different genera. and hence the pedigree method is not applicable to such cases. and potatoes. and both must have the superior character. Thus the value of an animal for breeding purposes depends on its pedigree. tobacco. The older method of breeding was to judge an animal entirely on the basis of appearance or performance. Hybridization.

" It is an attempt to apply to man the same principles of scientific breeding that have been so effectively used in the improvement of plants and animals. dogs. and so the characters of the hybrid can be preserved indefinitely. Many hybrids are more vigorous than either of thoir parents. Hybrid vigor may express itself in more rapid growth. Another value of hybridization is to increase vigor. Where sexual reproduction is necessary. hybrids may yield as much as 50 per cent more grain than the average yield of their parents. etc. either physically or mentally. medicine. In other words. not the race. in larger parts. If some of these represent what is desired. human . they seek to improve the surroundings. philanthropy. deal primarily with environmental factors. Man cannot be improved racially by improving the environment any more than a breeder can create prize winners from scrub stock by giving them the best of food and care. eugenics is an effort to control Most of the agencies evolution by selective mating. religion. Because segregation occurs when the hybrids are interbred. the situation is far more difficult. grows twice as rapidly as either parent and greatly exceeds them in height. it should be realized that they affect only the individual. Among animals. In corn. cattle. horses. or in other ways. Burbank's royal walnut/ a cross between the California walnut (Juglans californica) and the black walnut of New 7 England (Juglans nigra). concerned with the improvement of man education. " example. as in many plants and in all domesticated animals. and probably many others. EUGENICS The science of eugenics was founded by Francis Gal ton (18221911) and was defined by him as "the study of agencies under social control that may improve or impair the racial qualities of future generations. sheep. the members of the next generation are exceedingly diverse and represent all kinds of new character combinations. in For greater resistance to adverse conditions. selection being continued until a pure-breeding race is obtained. Although these influences are indispensable. they may be isolated and raised under pedigree culture.284 FUNDAMENTALS OF BIOLOGY without sexual reproduction there can be no segregation of genes. hybridization results in greater vigor in hogs.

Until recent years it has been rather generally assumed that environmental influences are of the greater importance.APPLICATION OF HEREDITARY PRINCIPLES Heredity 285 of and Environment. (2) Experimental breeding is not reasons for this. Environment merely gives an inherent tendency an opportunity to develop along lines predetermined by heredity. that is. and the results are less certain than those obtained with the lower animals. the surroundings determines the characteristics an individual comes to have are old ideas. The study of human heredity presents a number of difficulties. when fraternal twins. 260) are separated in early life and each is brought up under a different set of surroundings. it may never express itself. Our innate The environcapacities and tendencies are fixed by heredity. than any of the lower animals and subject to a much greater set of environmental influences. The study of twins furnishes strong evidence as to the greater importance of heredity in development. not a creative one. As Galton first showed. in the absence of an opportunity for sound training. Modern biology teaches that the more powerful influence in determining the constitution of an organism is heredity. but improvement itself depends upon the presence of favorable circumstances. Thus. when identical twins (see p. but that environment has an important part to play in the final expression of adult characteristics and qualities. On the other hand. Capacities for improvement are inherited. One may inherit exceptional musical ability. no amount of training can create a musician from a person with no natural aptitude in this direction. The relative* importance heredity and environment in the development of an individual has long been a matter of controversy. especially mentally. . what we shall do with our natural gifts. the environment is a guiding influence. ment merely determines how they shall develop. the dissimilarity between them does not diminish but usually increases as time goes on. Hereditary Characters in Man. and many people still have this That all men are born equal and that the nature of opinion. which always have a different set of hereditary characters. their resemblances in both mind and body persist. are brought up under exactly the same set of environmental influences. There are several (1) Man is more complex. On the other hand. but. for the most part. and others have later substantiated.

skin color. 3 Sex-linked (seep. A Some of these are: general ability. memory. The in most cases following characters are blending in their inheritance. offspring Although many human characters are known to be heritable. hair form (degree of curliness). but the mode of their inheritance is uncertain. literary mechanical . and arise from the action of several independently inheritable genes: general body size. great many characters are known to be largely determined by heredity. and all other shades but pure blue. shape of head. artistic mental ability. temperament. etc. and those concerning which we have the greatest amount of information are mostly abnormal or defective characters. stature. of many hereditary lines. incomplete and unreliable and even at best show only average conditions. weight. and consequently we must rely for information upon (3) Such data are often family records and other vital statistics. musical ability. (4) Every individual represents a complex mixture (5) The production of relatively few makes it difficult to obtain ratios indicating the chances involved and to determine what all the hereditary possibilities may be in any given mating. facial features. ability.286 FUNDAMENTALS OF BIOLOGY possible. much uncertainty exists regarding their mode of transmission. The case is not quite so simple. hazel. 2 Including green. A brief list is given below of some of the hereditary characters in man : DOMINANT Dark hair Brown eyes 2 Hereditary cataract RECESSIVE* Blonde hair Blue eyes Normal eyes Albinism Normal pigmentation Brachydactyly (short digits) Polydactyly (extra digits) Syndactyly (fused digits) Normal Normal Normal digits digits digits Normal Normal Normal Normal Normal Normal Normal Normal 1 Hereditary feeble-mindedness Hereditary epilepsy Hereditary insanity Congenital deaf-mutism Left-handedness Hemophilia (profuse bleeding) 3 Color blindness 3 Night blindness 3 In a few of the cases listed here it is a slight overstatement to say that one of the characters is recessive. 276).

and longevity. By defectives Although* many of these are confined to prisons. asylums. the intelligence of the individual remaining Some cases of feeble-mindedness may be caused by child-like. unu- The urgent need of eugenics is The highest birth rate exists among the classes sually high. Although . almshouses. baldness. but it has been found that at least 60 per cent of them are due to an inherited tendency and cannot be corrected. tendency to produce twins. it has been conservatively estimated that there are between 400. evidence that many forms of insanity are inherited. among the mentally inferior. It is the principal object of eugenics to discover the causes underlying this deplorable situation and to seek means of remedying it. Eugenics has two definite aims or pur- poses: (1) to eliminate undesirable qualities from the race by preventing the breeding of defectives (2) to increase the propor. a great many defectives are at large. it seems to be There is also ample inherited as a simple Mendelian recessive. The condition is a complex one. Although many defectives may be the victims of a poor environment and owe their unfortunate condition largely to this fact. mindedness results from a failure of the mind to continue its normal development. Although most of these people are not a direct menace to society individually. and similar institutions. as a class they are reproducing at a higher rate than normal persons. tion of superior strains in the general population by encouraging matings between highly endowed individuals. In both of those ways the average quality ing. beggars. of the race may be kept from deteriorat- apparent when it is realized that the present rate of reproduction among the mentally superior individuals is unusually low. allergy. Aims of Eugenics. is meant not only the feeble-minded and insane. For example. probably representing a combination of several defects.APPLICATION OF HEREDITARY PRINCIPLES 287 ability. accident or disease. having the least value to society.000 and 800. and all other persons who are a burden to society. astigmatism.000 feeble-minded persons in the United States of which perhaps only one-tenth are confined in institutions. For the most part. Elimination of Defectives. but criminals. there is no question but that most defectives are mentally Feebledeficient and simply lack the capacity for improvement. free to propagate their kind. paupers. tramps. nearsightedness.

the exact mode of its transmission is not understood.288 FUNDAMENTALS OF BIOLOGY insanity seems to be a recessive character. that is. This is due to the unreliability of the data. The same is true of paupers. to the fact that the condition may be greatly modified is by the environment. is tomy) but in the female a similar operation (salpingecattended with as much inconvenience and danger as any . The tendency to commit crimes is closely associated with feeble-mindedness. to be recessive. An attempt has been made to meet the situation by legislation. drunkards. etc. As a means of eliminating defective members of society. a number of plans have been suggested. Our present immigration laws keep defectives from entering the country but are inadequate because many immigrants who appear to be normal carry latent if It is evident that hereditary defects. and because the term variety of disordered nervous conditions. for the unfit. both heterozygous for the same defect. they may become their presence being suspected. as the percentage of illegitimate births among the socially worthless is very high. In the male this is accomplished by a very simple operation (vasectomy) performed under a local anesthetic and involving no risk to health. however. tives tend to intermarry. Mental " " tests performed on juvenile criminals in state reformatories have shown 50 to 90 per cent to be feeble-minded. many criminals being mentally defective. and " " except in extreme cases to grant them the same personal rights The result has been a constant multiplias normal individuals. however. to give them every opportunity to improve. used to cover a great Because most human defects. the inferior strains will eventually die out. For this reason. but many of these are not The tendency has always been to protect and care practical. may produce some abnormal children. arc heterozygous. have no effect. intermarriage among Defecfamilies having hereditary defects is to be condemned. because they are largely avoided by normal persons. Laws preventing the marriage of defectives. seem latent in many strains without Consequently two normal indi- viduals. both physical and mental. Laws compelling the segregation of the sexes in institutions are effective if enforced. defectives are prevented from leaving offspring. cation in numbers. but the most satisfactory plan is sterilization. but laws cannot be made effective except in a limited number of cases. prostitutes.

there In neither case. not to inherent infertility. so low in fact. The low birth rate of superiors is largely due to economic facLate marriage is unquestionably tors. Although sterilization is the only certain way of preventing the breeding of defectives. an important cause. etc.APPLICATION OF HEREDITARY PRINCIPLES other abdominal operation. this arises chiefly from the necessity of obtaining an education. as many people have no desire for children.690. It we know little concerning the manner of their has been pointed out that the birth rate among the intellectual classes is lower than among the population as a whole. public sentiment is not yet sufficiently enlightened as to the necessity of enacting and enforcing rigid sterilization laws. and the superior stocks do not average two. That superior mental ability runs in families was first proved by Galton and is accepted today as a Not only is general intellectual capacity inherited but fact. the total number of eugenic sterilizations performed in state institutions has been only 30. nearly half of which have taken place in California. For example. while luxury and selfishness must be added as a third factor. while but three-fourths of the graduates of men's colleges marry. but some seldom put them into operation. The high cost of maintaining modern standards of living is also a vital factor in reducing the number of children. Up to the end of 1938. literature. transmission. About 30 states have such laws at the present time. art. any interference with normal instincts or functions. that they are not maintaining themselves. No class can persist indefinitely unless there is an average of 3. Increase of Superiors. . it has been shown that only half of the graduates of women's colleges marry as compared with over 90 per cent of all women. however. 289 which the reproductive cells is In both operations the ducts through must pass are severed and sealed. except that reproduction is made impossible. The graduates of coeducational institutions average only 5 to 9 per cent higher.7 children per family. also special aptitudes for music. Although we that other fact these and the mental many recognize qualities are inherited.

an eye. owes its distinctive ular kinds of work. air. but to other organisms as well. exhibits a totally different form and structure because its functions are different. The former set of factors constitutes the inorganic environ- The branch of biology ment. various organs and the functions they perform. water. the organic environment. absorbs water therefrom. anchors the plant in the soil. that organisms are suited to complex of external conditions is under which they light. that is.CHAPTER XYII1 ADAPTATION In making a detailed study of any plant or animal. or of its structure. and environment is what ordinarily is implied by the term adaptation. structural differentiation has come about through a specialization of different parts of the body for particA root. 290 . the leaf being primarily fitted to carry on the work of photosynthesis and to regulate transpiraThe same correlation between structure and function is tion. on the other hand. is called ecology.. and carries it to the shoot system. The marvelous adjustment exhibited by all organisms between structure. the latter. It is the fitness of organisms to their conditions of life and the fitness of their parts to the functions they perform. function. One of the any other part of the body. features to the fact that it also everywhere apparent in the animal body. A leaf. The structure of the heart has significance only when we understand its mode of action. by reason most obvious facts in nature their surroundings to the great live. Not only is every organ. one is impressed with the striking correlation that exists between the structure of its As has been seen. tion exists between structure. a limb. that deals with the life relations of organisms. fitted to certain definite but a marked three-fold relaperform functions. function. organism is related not only to etc. and environment. for example. and the same is true of the stomach. with their relations to the conditions under which they live. An temperature.

the appearance of characters in an organism is particular set of hereditary factors in the dependent upon the presence of a chromatin of all of its cells. Tbus the plants and animals living in an alpine habitat are unlike lowland forms. prairies. and environment has been largely determined by the organism's ancestry. acters. the ones to trivial that often serve distinguish closely especially related species from one another. the environmental conditions in the two places are because chiefly For the same reason. The most important factors governing the distribution of plants into various kinds of habitats are chiefly those arising In animals the nature differences in soil and climate. Any set of conditions capable of supporting life constitutes a habitat. but most of the basic features of plants and animals do relation to the environment. forests. from and abundance of particular kinds of food are perhaps of major importance. each has its own distinctive fauna and flora. nearly body RACIAL ADAPTATIONS We are now ready racial adaptations cases where to consider a few conspicuous examples of an intimate relation between structure. show a cance. swamps. deserts Ponds. seashores. The term " adaptation" is applied not only to the fact of fit- Some characters. like most other charThus heredity. in the deep sea. These arise. through the great internal influence inborn in are the that all adaptations racial. and in any given habitat only such plants and animals can live as are adapted in structure and function to meet the particular conditions that characterize it. but to adaptive characters themselves. rivers. In general. meadows. tide-pool animals are not found dissimilar. is. have no obvious adaptive signifiness. organism. although physical factors also play a prominent role. and these are largely the nature of the local conditions.ADAPTATION The structural differences displayed by organisms living under diverse sets of external conditions are largely correlated with the conditions themselves. for the most be may the surroundings themselves do not cause an organism to part develop adaptive features. and although the final expression of many characters influenced by certain environmental factors. Since every individual has . function. This arises from the fact that organisms which occur in the same habitat have similar fulfilled by life requirements. or the plants of a moist meadow upon the desert.

A thirk-loaved desert plant (Agave deserti).3.292 FVNDAMENTALK OF BIOLOGY '' ' in FIG. . 10. the fleshy leaves serving as reservoirs for the accumulation of water.

and this is solved by desert plants both by increasing absorption and retarddry air. no leaves being present.. viz." many more examples will be suggested by the few that are given. A barrel cactus (Echmocwtus cylmdraccus). live there as are structurally . high temperatures. Desert Plants. dry soil. Plants that live in deserts exist under a set of very severe conditions. ( Only such plants can adapted to endure these rigorous conditions (Fig. It is evident that the greatest problem of the desert environment is the conservation of water. intense light. thickFTC. 192). longitudinally fluted stem stores water and carries on photosynthesis. and often strong winds. 1 K ened.ADAPTATION 293 " been called a bundle of racial adaptations. The groon.

at the same time. as in the cacti (Figs. hinders photosynthesis. Another adaptation seen in many desert plants is the development of water-storage tissue. In most desert plants the FIG. leaves are small. Cross section of a leaf of oleander (Nerium'). Various structural adaptations bring about the same results in different cases. 193 and 194). As a compensation. either in the leaves. many small-leaved and almost leafless desert plants have green stems. live in cells. Reduction of leaf surface effectively lessens transpiration but. as the following examples show. root system but a very extensive one. as in the cacti. Many desert them plants have a very deep root system that enables Others have a shallow to absorb water from a great depth. the several layers of epidermal upper and lower palisade tissue. a plant adapted to dry air.294 FUNDAMENTALS OF BIOLOGY ing transpiration. or in the stems. 195. often being developed as mere scales or spines. Many plants of dry regions have hard stiff leaves with an . and stomata in pits protected by epidermal hairs. Note ths thick cuticle. X 200. as in the aloe and agave.

under are ordinary garden conditions. These plants branches the plant Animals carry seeds in several different ways. dry. one-seeded fruits. in some cases causing the seed to spin around and thus retard its descent. they retain regions grown their desert characteristics. Desert plants are peculiar because they live under a set of extraordinary conditions. A strong wind might carry such a seed a considerable distance from the parent plant. characteristic of many desert plants. the two most important ones being wind and animals. 197). are eaten in 1 great numbers by various kinds of birds and mammals. ash. 195). Trees such as the elm. The dandelion. a tuft of cotton or down acts as a parachute and keeps them suspended in the air for a long time. The seeds " of the various kinds of tumble weeds. Many marsh plants owe their wide distribution to the fact that their seeds are carried in the mud that Fleshy fruits. Seed Dispersal. small. brightly colored ones and those attractively flavored." such as the Russian thistle. and for the are structurally adjusted to most part their distinctive Thus when most plants of dry features are racial. 196). not acquired. dropping its seeds as it moves along. They meet these conditions. 1 maple. When the of the way. Such seeds are often carried great distances by the wind. by the wind in another grow on open assumes a spherical seeds are ripe the entire plant breaks off at the ground and is blown about by the wind. and various kinds of "stick-tights" serve as a means of attachment to the bodies of animals (Fig. the "seeds" are really is of no significance here. . The barbed appendages of the cocklebur. thistle.ADAPTATION 295 unusually thick cuticle. of a thick covering of hairs. cottonwood. and catalpa have winged seeds adapted to dispersal by In falling to the ground the wing presents the wind (Fig. are dispersed plains. burdock. but this distinction la some of these and the following examples. which acts as a check against transpiraIn such cases the stomata are generally sunken below the tion. By an incurving form. is also thought to be related to the conservation of Hairs are outgrowths from the epidermal cells. The seeds of many plants are adapted to be disseminated by some particular agency. milkweed. The presence water. level of the leaf surface and often are confined to pits (Fig. particularly clings to the feet of wading birds. a flat surface to the air. and many other plants have downy seeds.

ticks. burdock. winged fruit of ash. Insect Pollination. D Flowers are either self-pollinated or crosspollinated. C and />. winged fruit of box elder. X 2. downy seed of milkweed. but in many cases when swallowed they pass through the body unaffected by the digestive fluids. winged seed of catalpa. 196. Cross-pollination is brought about chiefly. B. depending upon whether each flower pollinates itself or whether pollen Jfrom one flower is transferred to the pistil of another flower. horehound. cocklebur. (C. by wind or by Self-pollinated flowers and those cross- .296 FUNDAMENTALS OF BIOLOGY Often the seeds are voided without being swallowed. D. F. C. B. downy fruit of dandelion. (A and B. winged fruit of elm. A. E.) ARC insects.) FIG. beggar. A. Seed dispersal by wind. 197. Seed dispersal by animals. X 1. others. Flo. C. />. XI. X 2.

are largely dependent upon butterflies or moths for their polli- . the act of pollination being purely an is incidental matter. some part of the insect's body comes in contact with the stamens and FIG. 200). 297 and odor- flowers chiefly to get nectar from them. pink. The stamens are situated at the mouth of the corolla. (From Sanderson and Jackson. pollination is be understood accomplished. 61) or in a special sac or spur (Fig." Oinn and Company. It unavoidably carries pollen from flower to flower in search Butterflies their and moths obtain nectar for own fed by use. etc. 198). is covered with pollen. In get- ting nectar from a flower.ADAPTATION pollinated by the wind are usually inconspicuous while most flowers crossless. to be pollinated by butterflies or moths. a sweet liquid secreted by glands situated inside the flower. showing coiled proboscis. petunia. as the caterpillars are not the adults. have a tubular or funnel-shaped corolla at the bottom of which nectar collects (Fig. that there It is should nothing intentional in this behavior on the part of the insect. Side view head of butterfly. Such flowers coiled (Fig. by permission. under the head like a watch spring Flowers that are adapted 199). Flower of garden nasturtium (Tropaeolum) cut through the middle to show the spur (a) and the nectary (n). natural size.. either at the base of the petals (Fig. "Elementary Entomology. which when not in use is of 199. Insects gather Nectar nectar as food. This is then carried to the next flower where some is rubbed In this way crossoff on the stigma. The mouth apparatus of a butterfly consists of a very long tube called a proboscis. pollinated by insects have white or brightly colored corollas or an attractive fragrance Insects visit . 198. either for themselves or for the larvae. such as the tobacco. in going of food. honeysuckle.) FIG.

it comes in immediate contact with the stigma.") by bees (Fig. off on the stigmas of other flowers. (From Snodgrass. Flower Petunia. by pollination Nectar collects at the bottom of the tubular corolla. thus rubbing off some of the pollen brought of lower lip or platform of its body causing the corolla to open. violet. "Anatomy of the Honeybee. It then encounters the stamens. which often In all pollinators. Although this pollen is lost to the plant. Bees gather both nectar and pollen. the weight The pistil is so situated that. a type adapted to moths. sweet pea. Left hind leg of honeybee (worker).298 FUNDAMENTALS OF BIOLOGY nation. which they feed to the In many bees a portion of the hind legs serves as a " " in which pollen is carried to the hive and fed pollen basket to the larvae (Fig. belonging to different families. 202). and sage are flowers especially adapted for pollination of FIG. as the bee enters the flower. showing pollen basket (Cb) on outer surface of tibia (Tb). that which adheres to other parts of the body may be rubbed The snapdragon. Normally the flower is more or less closed. 201). in fact Although these flowers are not related they have a highly specialized corolla somewhat similarly modified. closure of the corolla makes the essential organs inaccessible to ants. larvae. steal pollen but are practically of no value as the flowers mentioned above there is a sort upon which a bee may alight. 201. as these insects are particularly adapted to reach the nectar. FIG. from another flower. 200. usually . the parts of the corolla fitting The together in such a way that the stamons and pistil are hidden.

and thereby acquires a gathers the nectar and departs. D. 202. it body harmonizes with the surroundings. 203). color. A. C. thus inconspicuous and affording concealment from A elongated body and legs shows a striking resemblance to a branched twig. snapdragon. but a few will out the general situation (Fig. and even the worm- . veining. and thus is very difficult to see. B D many animals the color Flowers pollinated by boos. redrawn after Avebury. The walking-stick insect with its slender great suffice to point rendering enemies. (A. and when disturbed some of them assume a rigid position at an acute angle from the stem. The measuring worms. been removed. When its wings are folded. The dead-leaf butterfly (Kallima) of India is one of the most amazing instances of protective resemblance known. In C the right lateral petal has corolla. showing bee entering FIG. which are caterpillars of certain moths. violet. or form of the In a great many examples could be given. new lot of pollen. B. in regard to the shape. petiole. The green body of the katydid with its veined wings blends with the foliage upon which it feeds. common sage. it exhibits almost perfect similarity to an attached dead leaf.ADAPTATION located below the Finally it 299 pistil.) Protective Resemblance. sweet pea. are greatly like twigs in form and color.

or vice versa. katydid. walkingon a twig. such as the . Prairie and desert forms. their upper holes (Fig.. larva of geometric! moth resting extended from a twig.} The color change of the chameleon and certain other lizards from brown to green. 204). by permission. 203. Fia. B. stick insect Protective resemblance among insects. from Riley. New York.300 FUNDAMENTALS OF BIOLOGY When the wings are outspread. Inc. Appleton-Century Company. to harmonize with the background is well known. surfaces are seen to be brightly colored. D. "Evolution and Animal Life" (A. A protective coloration of mammals in the wild state is very common. C. A. B and C\ from Jordan and Kellogg.

Arctic fox. and weasel being good examples. at rest on a leafy twig. 301 and lion. crack nuts. Some of these forms are colored not so much striped. at least during the winter. 204. pick fruit. the bill . with wings extended. B. and leopard tend to be spotted. kill smaller birds. A generalized bird. Arctic animals are usually white. One of the most striking illustrations performance of diverse functions. and break open eggs.-The dead-leaf butterfly (KalUma) of India. and of adaptations in animals bills of birds for the is seen in the modification of the feet insects. uses its bill for many different purposes. It can capture 205). as in many similar cases. As a consequence of this varied diet. or mottled. an excellent example of which is the crow (Fig. which they feed. for protection but to render them inconspicuous to animals upon FIG. the polar bear.. such as the zebra. one-half natural size. tiger.ADAPTATION wolf. dig corn. tend to be brown or gray. camel. the protective value of their coloration has been greatly overruled Feet and Bills of Birds. while in others. Forest-dwelling types.

feed mostly on seeds. poorly developed feet. A few examples will be given. Insect-eating birds are of several different types. They . lengthwise on a limb or on the ground. stout. etc. Similarly its feet are general- being used for walking. grouse. FUNDAMENTALS OF BIOLOGY crow is in no way modified. a generalized bird. Their bills are straight. short toes (Fig. Hawks.302 of the ized. 2064). strong. X H- ground. the latter acting as a brace against the tree. They short. 206C). and chisel-like. walkThe more restricted a bird's diet and the ing on snow. scratching. some being more highly specialized than others. turkeys. scratching birds are in general poor fliers. 205. These birds have very small bills surrounded by bristles. more specialized its method of getting food. the greater is the amount of structural modification that it exhibits. 206D). stout. Crow (Corvua brachyrhynchos) .. Scratching birds. perching. in consequence of which their bills are and curved. eagles. etc. 206B). the body ivsf ing. such as chickens. wading. Spending most of their time on the FIG. quail. and other birds of prey have powerful feet provided with sharp curved claws adapted to seize and carry living prey. and strong hooked beaks for tearing flesh (Fig. The nighthawks and whippoorwills are night feeders that capture insects on the wing (Fig. have short legs arid straight. the tongue sharp pointed and barbed. The feet are sharp clawed and the tail feathers stiff. The woodpeckers dig into tree trunks for grubs and beetles (Fig. They have long narrow wings and very The latter are too weak to be used in walking or even to support the body but are used merely for clinging.

E. D . X Y\ D. a woodpecker. B. They are the smallest of birds. fly 303 swallowing C . 206. their rapidly moving wings holding the body poised in front of the flower without coming to rest. X J^. The hummingbirds . A. red-shafted flicker (Colaptes cafer). California quail (Lophortyx californica) a scratching bird. Five types of land birds showing adaptation of feet and bills to FIG. a nectar feeder. a night-flying insectivorous bird. poorwill (Phalaenoptilus nuttalli). C. X ^.ADAPTATION through the air with the mouth widely open such insects as happen to be in the way. their long slender bills being well fitted for probing into flowers (Fig. X Y*>'. pigeon hawk (Falco columbarius) a bird of prey. X M are nectar feeders. hummingbird (Cdlypte anna). 2061?). diverse life habits.

black-neoked stilt B. green. the mud. a diving bird. bird. in either case preventing the bird from sinking into B A. while in the stilts. or shorter and more or less webbed. C't pied-billed grebe (PodUymbus podiceps). X M. sandpipers. and plovers the bill is long but slender and . The toes are either very long and thin. X /^. a wading . 207 A). X }. The cranes and herons have long. sharp-pointed beaks for spearing fish. lift the body above the water. (Himantopus mexicanus). of Water birds three different types. Because the legs neck and bill snipes. the are elongated to permit feeding in the water or in the mud on the bottom.304 FUNDAMENTALS OF BIOLOGY Wading birds are characterized by long slender legs (Fig.winged teal (Nettion carolinense) a swimming bird of the duck type. stout. 207. FIG.

and a general mouse-like appearance Shrews live chiefly on insects. These can be studied to good advantage in man or in any of the monkeys. short legs. For example. In most cases. 2 canines. a dog has 42 teeth. along shore or in shallow water. In typical mammals there are four distinct kinds of teeth: incisors. small eyes. pentadactyl limbs. and walking ^yith the entire foot resting flat upon the ground. X the mammalian body. where all cialized condition (Fig. The sharp-pointed beak is well fitted for grasping fishes. 209). four types are present in a relatively unspeThe primitive number of teeth in mammals is 44. etc. 208. 2075). as in all other animal groups. Among the most primitive of placental mamjnals are the shrews small. Grebes and loons are diving birds. and but slightly differentiated. swimming under water both to capture fishes and to avoid their enemies (Fig. this number has been reduced. and molars. a relatively unspecialized mammal. 8 premolars. Although structural modification may affect any part of FIG. two of the . their teeth being small. mostly terrestrial forms with pointed heads. broad pouched and flat ducks and geese. canines. there being in each jaw 6 incisors. having relatively short. a striking correlation between structure and life habits is seen. it is particularly evident in teeth and limbs. and arc usually awkward on land (Fig. being used to probe for small worms. crustaceans. etc. being pelicans. In mammals. strong and hooked in the gulls. 208). and the feet placed far back on the body. Generalized Mammals. sharp-pointed. premolars. and 6 molars. (Fig. Teeth of Mammals. mollusks. Their bills are modified in the in tho in accordance with their feeding habits. however. a type of progression known as plantigrade. A shrew (Sorex ornatus). so that on land an erect position is assumed. 207C).ADAPTATION more or less soft at 305 the tip. Swimming birds have lobed or webbed feet. The toes are webbed or lobed. The most generalized mammals are nearly all ground dwellers.

ing. while the premolars and molars arc flat-crowned and adapted for grindThe total number of teeth is 20. pairs of incisors. adapted for gnawing. canines are absent. mice. but the number of other teeth is the same as in man. X }.. In the rodents. A tendency in man to reduce the number still further is seen in the late appearance and often imperfect development of the third molars ("wisdom teeth"). In man 32 teeth are present. and 6 molars. guinea pigs. making a total of 36. Skull of spider monkey (Ateles). . beavers. etc. a mammal whose teeth are relaThree pairs of premolars are present in each jaw. are large and chisel-like. 2 canines. squirrels. 209.306 FUNDAMENTALS OF BIOLOGY upper molars being absent. The two Fio. tively unspecialized. rabbits. 4 premolars. a herbivorous group including the rats. the teeth . Skull of beaver (Castor canadensis) a rodent. 210. gophers. Fio. X J^. there being in each jaw 4 incisors.

making a total of 20 or 22 teeth (Fig. 211. No canines are ever present. are is only 16. 210). teeth are present in the male. The skunks. The total number is The incisors Skull of horse (Equus caballus). The teeth of hares and rabbits are more numerous (totalling 30) and less highly specialized than those of other rodents. an ungulate. FIG. foxes. cats. FIG. The total number of teeth is 40. of which 4 are usually present. Small canine are adapted for cropping. badgers. The incisors. but absent in the female. teeth are adapted for cutting and tearing flesh. but some The 42. 211). X K. raccoons. The cats have the lowest . while the 12 molars are flat crowned and used for grinding. bears. carnivores include the wolves. X /. 212. very large and chisel-like. In most cases the teeth are adapted for etc. in many cases. tearing and cutting flesh (Fig.ADAPTATION 307 are adapted for gnawing. and the total number. rodents have 4 or even 6 premolars. Skull of timber wolf (Canis occidentalism a carnivore. the premolars and molars for grinding.



(30) and the most highly specialized teeth of this type, the bears (with 42 teeth) the least specialized. The incisors are small and of little use, the canines are large and pointed, while the premolars and molars are sharply ridged.



dentition of the ungulates, or hoofed




highly specialized for a herbivorous diet. To this group belong the cattle, sheep, goats, deer, hogs, horses, etc. Hogs have the complete set of 44 teeth, the primitive number. Horses

have 40 teeth, there being only 6 premolars in each jaw (Fig. 212). In most of the other ungulates all four kinds of teeth are present, but the total number is still further reduced. The incisors, which are used for cropping grass or herbage, are generally large and chisel-like, while the molars and premolars are broad and flat and provided with a complex surface for grinding. The deer, cattle, sheep, and other ruminants have 28, 32, or 34 teeth; there are no upper incisors, the lower incisors biting against a horny pad. Here the upper canines are absent, the lower canines are incisor-like, while the number of premolars in each jaw has been reduced to 6, as in horses.

One order



the edentates,


characterized by

an imperfect development or total absence of teeth. To this group belong the sloths, armadillos, and most of the anteaters. A
total absence of teeth

also characteristic of
It will


of the whales.

be recalled that, in a typical of an upper arm, forearm, fore limb consists the vertebrate, the limb a thigh, shank, ankle, and hind of and hand wrist, foot. In both cases the primitive number of digits is five, In a typical mammal the the limb thus being pentadactyl. the of a arm consists single upper bone, humerus, and the forearm The wrist is made up of a of two bones, the radius and ulna. number of small bones called carpals. These are followed by the



forming the palm of the hand, and the five sets The corresponding bones in the hind limb are the femur, forming the thigh, and the tibia and The ankle consists of a number of fibula, forming the shank. small tarsals, and the foot of five metatarsals, which form the sole,
five metacarpals,

of phalanges or finger bones.

phalanges or toe bones. the fore and hind limbs of mammals are constructed Although to the general plan described above, in most groups according
five sets of


differences in detail

have arisen in accordance with diverse func-

tions (Figs. 213


and 214). Specialization of the limbs of mammals has involved changes in the form and relative length of the bones, a reduction in their number, and often some degree of fusion between certain bones. There are five chief classes of functions for which mammalian These are: (1) limbs, in different cases, have become modified.

Fro. 213.



Right fore limbs of rhesus monkey, A; wolf, B\ and horse, 6'; h, humerus; r, radius; u, ulna; c, carpals; w, metacarpals; p, phalanges; / to V, The phalanges are the individual bones comprising the digits. digits.
fossorial, for digging

and burrowing;

(2) cursorial, for

rapid runconsider

ning on the ground;

(3) arboreal, for living in trees: (4) natatorial,



for flying.




specific instances of

each type of functional adaptation.

Fossorial Adaptation. number of mammals, especially many of the rodents, excavate burrows and live in them but in most The moles, cases they spend much of their time at the surface.


however, live entirely underground, and consequently are very highly specialized for a fossorial existence (Figs. 214J5 and 215).



The spindle-shaped body with
narrow shoulders, and short
feet being provided



pointed sncut, short neck, obviously adaptive. The

extremely short limbs are modified for digging, the hands and with long strong claws. External ears are



Fia. 214.

Right fore limbs of mastiff bat, A; mole, B\ and harbor
Labeling as in Fig. 213.


FIG. 215.


mole (Scapanus latimanus), a


mammal, X


Moles are insectivores, belongabsent, and the eyes are vestigial. ing to the same order of mammals as the shrews and hedgehogs. Their teeth are simple.
Cursorial Adaptation. In mammals that are adapted to run rapidly over the ground, the body is relatively light and the limbs long and slender. The wolves and foxes have limbs pro-



vided with four or five clawed digits. Elongation of the arms and legs has been brought about by an elevation of the wrist and heel above the ground, so that these animals walk on their toe* This method of progression is designated digiti(Fig. 2135). In most grade. ungulates, on the other hand, the digits have

been reduced to either three, two, or one, the others being functionless or entirely absent. Ungulates typically walk on the tips of their toes, the hoof corresponding to the nail or claw of othei mammals (Fig. 213C). Animals that walk in this manner are said to be unguligrade. Horses and deer are good examples oi unguligrade runners. In many cursorial mammals the radiu? and ulna are fused together, as well as the tibia and fibula. Thi? prevents a rotation of the limb but enables it to move very effectively in one plane.

FIG. 216.



(Phoca vitulina), adaptation,








mammal adapted

to living in trees.

The monkeys represent a type of Both the fore and hind

limbs are modified for grasping branches, the first digit being opposable to the others (Fig. 2134). The tail is long, and in the
is prehensile, thus being of great utility swings from branch to branch. Together with the apes and man, the monkeys form a group of their own, Even the few ground-dwelling primates show the primates. many anatomical evidences, especially in the embryo, of a former arboreal existence. Other arboreal mammals are the

New World


to the animal as

squirrels and the sloths, the former running along branches, the latter suspending themselves beneath them.


The seals are aquatic mammals Natatorial Adaptation. belonging to the carnivore group (Figs. 214C and 216). They are not so highly specialized for natatorial existence as the whales and their relatives (cetaceans) or as the manatees and dugongs







of the seal with its



smooth fur

obviously adapted to
are paddle-like, the

progression through the water.

The limbs

upper portion being short, but the hands and feet are elongated and provided with completely webbed digits. The related sea lions and walruses can use their hind limbs in walking, but the
seals cannot, thus being

Volant Adaptation.
flying squirrels,

more highly adapted to aquatic life. Although some mammals, such as the are adapted for gliding through the air for short

distances, the only mammals that really fly are the bats (Figs. 214,4 and 217). To serve this function their bodies are highly The forearm and four of the digits are greatly modified.

elongated; these support a thin fold of skin that reaches to the body and to the hind limbs, forming a wing. The thumb is

FIG. 217.


bat (Lasionycteris nocti vagans), volant adaptation, X 8 6-


mammal showing


great difficulty in

Bats have shorter than the other digits and is clawed. moving on the ground. When at rest they
of their

hang downward from a support by means



or with head up, suspending themselves by their wing-claws. As in birds the bones are light and the pectoral muscles highly


Adaptive Radiation. The feet and bills of mammals, and the limbs of mammals furnish

birds, the teeth of

of the law of adaptive radiation, which is widely divergent forms in animals ancestrally of the

excellent examples "the development of

same stock

or of related stocks, as a result of bodily adaptation to widely

environments (Osborn). Originally all the members of each group were primitive, generalized forms living under the same conditions, restricted in distribution, with similar life




As migrations into habits, and hence structurally much alike. different habitats took place, new conditions were encountered, and structural modification followed. Thus all mammals,
although fundamentally of similar structure due to descent from a common ancestry, show many differences as a result of adaptation to diverse conditions of living.

Convergent Adaptation.
different conditions, tend

Although related forms, living under toward divergence in form, as stated

FIG. 218. vertebrates.
poise (a

Convergent adaptation of form in three wholly unrelated marine A, shark (a fish); B, an ichthyosaur (an. extinct reptile); C, a por(After Osburn.}


above, unrelated forms, living under similar conditions, tend toward convergence of form. This is the law of convergent
adaptation, which has been stated as "the production of externally similar forms in adaptation to similar natural forces" (Osborn). Because the conditions for existence in the water require the
sort of adaptive features, we find a marked external resemblance between such wholly unrelated vertebrates as a shark (a fish), an ichthyosaur (an extinct reptile), and a porpoise Each has a spindle-shaped body, paddle-like (a mammal). anterior appendages, a dorsal median fin, and a somewhat



type of




Fundamentally, of course,



these animals are very different, since they have progressed along independent lines of descent.

The law


convergent adaptation


well illustrated in the case

of desert plants. Many species having the same general aspect to different families, the superficial resemblance belong widely

being due to their living under the same environmental conditions.

Although the structure of organisms and the way in which their parts function are determined mainly by heredity, the environment, in providing an opportunity for inherent tendencies
to express themselves, constantly exerts a modifying influence. It is apparent that in every habitat conditions constantly fluctuate, and that all organisms, by virtue of the power of

respond to these slight changes, mostly in ways advantageous to themselves. When extraordinary changes occur, however, as when an organism is removed from its natural environment and placed under a new set of conditions, it also may be able to adjust itself, especially if the change is made

very gradually. Adaptive Response. It is important to distinguish between the state of being adapted and the process of becoming adapted. An organism is adapted to the normal conditions of its particular
habitat chiefly

by reason

of its hereditary constitution,

and so

most adaptive characters are

To a


react favorably to


limited extent, howconditions that arise; in

other words,


become adapted to a new environment.
of adaptive response.



as the



reaction to

the changed conditions may be functional, structural, or both. Adaptive characters that thus arise as a direct response to external influences are spoken of as acquired (or individual)

A few examples will be given. Everyone knows that the growth of a plant

enced by temperature,
diseases, injuries, etc.

is greatly influmoisture, character of the soil, is


a plant


in darkness


to develop chlorophyll, its leaves remain small,

and the

stem becomes very long and weak (Fig. 219). Such shoots, arising from seeds deeply buried in the ground, are stimulated to reach up toward the light, where the leaves may enlarge and turn green. There are many other responses in plants where

clearly indicated.

When grown

in the shade,



kinds of plants have larger leaves than when grown bright sunlight, and so a greater amount of green tissue

exposed for carrying on p]n;io-\ MJ]I< -i-. Many plants develop more extensive root systems in dry soil than in moist The cuticle soil, thus enabling sufficient water to be absorbed.

on many leaves becomes thicker when they are grown under dry conditions, transpiration being thereby diminished. In the water buttercup and mermaid weed, the same plant produces finely dissected leaves beneath the water and entire leaves when


Two pots of bean seedlings exactly of the same ago. A, plants FIG. 219. In the latter the stems in the light; B, plants giown in total darkness. are white and the leaves pale yellow. Both sets of seedlings were given the same amount of water and kept at the same temperature.
grows up into the


Such plants exhibit extreme


in their response to external conditions.

Animals are influenced by temperature, moisture, the character and amount of food, exercise, diseases, injuries, etc., and many
brought about are adaptive. A familiar human skin through continuous exposure to bright sunlight. The formation of brown pigment, which in the negro is racial, tends to prevent further burning and consequent injury to the underlying tissues. The development of callosities on the skin through friction is also an adaptive
of the changes thus



the tanning of the

response, as






of muscles


persistent exercise.

Recovery from a disease often brings about a condition of immu-




lity to further attacks,
as caffeine


and a tolerance for certain poisons, such or nicotine, may be gradually acquired. important limitation that should be kept in mind regarding
of adaptive response


that only minor adjustments




basic features of organisms are incapable By a manipulation of the environment we

human stature, change eye color, increase potential intellectual ability, or alter any of the other In the characters that are determined largely by heredity. yast. majority of cases the amount of modification that may be
cannot increase individual

through a radical change in external influences is slight and always limited. The striking cases ordinarily mentioned are exceptional, some organisms being much more plastic than others and hence more susceptible to modification. Explanation of Adaptation. In speaking of the power of

" adapts adaptive response, it is often said that an organism This and itself" to this or that condition of the environment.

be avoided, as their

similar phrases implying conscious efforts or purpose should meaning is entirely metaphorical even in the
of intelligent animals.


reactions are advantageous,

while others are not, but in neither case can desire on the part of the organism produce a modification in structure or function.

For example, as waste products of metabolic activity, some plants produce substances that may happen to be bitter, poisonous, or otherwise unattractive to certain animals, and such plants may
be avoided for this reason.


to say that they produce sub-

stances "to protect themselves" from being eaten
entirely misleading in that

by animals


implies foresight on the part of the

adaptation is not purposive, how has it other biological problems, this is a never been satisfactorily answered. Most tions are in the nature of direct responses to

come about?



question that has individual adaptaexternal conditions,


we cannot


why an organism reacts to a given stimulus

Regarding the origin of racial adaptathey have arisen by a process of evolution, but as to how they have evolved there is much uncertainty. Two plausible theories have been suggested: one called the
tions, it is certain that

a characteristic way.

inheritance of acquired characters, the other, natural seleaLion. These are discussed in their proper setting in Chap. XXII.

but from organic compounds occurring in dead plants and animals or in the waste products of living organisms.CHAPTER XIX SAPROPHYTISM. para- of particular interest to the student of because illustrate some of the more striking biology they general ways in which organisms arc adapted to the living environment. and the breaking down of protein material. however. putrefaction. saprophytes down these complex organic compounds into simpler products and thus are the direct cause of decay. Ordinarily there is a succession of various kinds of decay-producing organisms. In the course of obtaining food for themselves. AND SYMBIOSIS It has been pointed out in the last chapter that organisms arc related in their life habits not only to the physical factors of their environment but to other organisms as well. Although the breaking down of caris called fermentation. chiefly 317 . we can give attention only to a few phenomena of saprophytism. Their supply of carbon comes not from the carbon dioxide of the air. organic acids. etc. such as various alcohols. these processes are essentially similar. and symbiosis are SAPROPHYTISM Saprophytes are plants without chlorophyll that absorb their food directly from dead organic matter. It is important to realize that the decomposition of all dead organic gradually break matter takes place entirely through the agency of living organisms. sitism. The decomposition of dead organic bohydrates matter involves a complicated series of changes. Each kind of none can living thing has reciprocal relations with other kinc^ The limitless ways in which organisms are live unto itself. that here The of its many aspects. which thereby obtain material and energy necessary to their own metabolism. PARASITISM. there usually being a large number of intermediate products formed. as does that of green plants. associated and interact with one another constitute such a vast subject. Decay of Organic Matter.

other substances can be utilized only after being acted upon by certain other bacteria that oxidize them in order to obtain a source of energy for themselves. tion are chiefly water (H 2 0). but the. methane (CH 4 ). have the power of forming nitrates directly from free nitrogen (see pp. ammonia (NHs). hydrogen sulphide. hauatorium. The water and carbon may be directly FNJ. of ing on the first bacteria oxidize methane. Mycelium of bread mold (Rhizopus) Riving rise to sporangia. while still converted to nitrites and then to nitrates which there are two types.318 FUNDAMENTALS OF BIOLOGY bacteria. sporangiophore. D. sporangium. /?. 332-333 and 359). and free hydrogen. (From Sinnott. Other kinds of is others. . "Botany. ammonia by the nitrifying bacteria.4. . horizontal branch. and free nitrogen. forming substances directly utilizable by green plants. carbon dioxide (CO 2 ). one carrychange. hydrogen sulphide (H 2 S). For example. the nitrogen-fixing and Problems. PrinciiH. but none except The ultimate products of decomposithe last ones exhausts it. All these organisms gain their subsistence from the great amount of potential energy contained in the dead material."} used again by green plants in the synthesis of new organic compounds. each carrying the process a little farther. 220. dioxide free hydrogen. (7. the other the second. until finally only a few relatively simple substances are left.

221. moist woods. pinesap (M. hypopitys). On the Indian pipe (Monotropa uniflora). the other fungi have a delicate. PARASITISM. Only a relatively few saprophytes occur among the seed plants. These plants dead animals. white. many others.SAPROPHYTISM. thread-like body called a mycelium (Fig. one of the best known being the Indian pipe (Monotropa . 27) is really only a spore-producing structure. examples being yeasts. molds. rotting yeasts are unicellular. on the right. AND SYMBIOSIS 319 are found Besides the bacteria of decay. mush- rooms. and logs. 220). the vegetative body of the plant being a mycelium that grows in the soil or in decaying wood. Two saprophytic seed plants found in rich. many other kinds of saprophytes among the fungi. The fleshy "mushroom" (Fig. etc. The live on humus. (Photograph supplied by McFarland Publicity Service.) Fia. but most of left.

absorbing its nourishment from the partially decayed organic matter contained in the soil. Diagram representing the circulation of elements through the air. often reaching a height of a foot. sulphur. for animals. calcium. "Botany.320 uniflora. All the other elements that become part of the plant body. FUNDAMENTALS OF BIOLOGY Fig. and for dependIt will be recalled that green plants obtain the entire supply of carbon. 222. magnesium. bears a few reduced loaves and a single terminal flower. The whole plant is white or sometimes The unbranched entirely without chlorophyll.. is 221). (From Sinnott. stem. The Indian pipe grows in rich moist reddish and woods. are absorbed as salts dissolved in the soil water (Fig. Sulpnur Calcium Magnesium Potassium LIrpn from x Rock Particles By Solution FIG. potassium. green plants provide a source of Formation of Carbohydrates Fats Proteins and Protoplasm f Nitrate I i Salte Phosphorus * I j. and that these elements enter into the formation of all the organic substances in plants and animals. iron. By virtue of their ability to construct food from inorganic substances. "--- . and bodies of organisms. Principles and Problems. etc. however. 222). such as nitrogen. Cycle of Elements. and oxygen that they use in food manufacture from the carbon dioxide of the air and the water of the soil.") ent plants. hydrogen. soil. phosphorus. material and energy for themselves.

waste products resulting from the breaking down of organic matter in respiration are constantly given off. So far as plants are concerned. Without ultimately return elements to the inorganic world. that saprophytes play a much more important role in the economy of nature than is commonly realized. and so have the same relation to their food supply as do dependent among animals that can live only one . During life. AND SYMBIOSIS It should be 321 understood that the chemical elements of which the various components of protoplasm and its derivatives are constructed are only temporarily associated in an organism. A careful study of Fig. All animals eat either dead or living organisms -(or their products). and because green plants supply all the food that other organisms utilize. In fact. PARASITISM is no sharp line of distincand parasitism. Eventually the carbon dioxide of the air and the mineral salts of the soil would become so It is evident. are either immediately available for use again by green plants or arc ultimately rendered so. in effecting this decomposition. them dead plants and animals would rapidly accumulate. All dependent plants absorb organic substances from an external source. 222 should make this situation clear.SAPROPHYTISM. many fungi may live either as saprophytes or as parathat is. but eventually all organic matter constructed by green plants is decomposed into its constituent elements. PARASITISM. and it makes little difference whether . and the various elements essential for life would be "locked up" and so rendered unavailable for green plants. in contrast to the obligate forms is way or the other.the source is living or dead. and then back to the inorganic world. there tion between saprophytism sites. both during life and afterward. A plant may be eaten by an animal and some of its substance become temporarily transformed into animal protoplasm. while after death destructive processes continue under the influence of other organisms. It is apparent that saprophytes. Because of all the products resulting from the breaking down protoplasm and its derivatives. there exists in nature a continual circulation of elements in its complete form. therefore. Similarly parasitism not a clearly defined phenomenon. to saprophytes. are facultative. from the inorganic world through green plants to animals. greatly depleted that life would cease.

being used to describe cases where one kind of organism lives in or on another. obtaining its food at the other's We The two organisms involved in this type of association expense. but the host is parasite always more or less injured. Parasitism. or a herbivorous animal and the vegetation upon which it feeds. benefits of the the all receive association. by the destruction of host tissue. the liberation of poisonous material (toxins). Not only docs the are known respectively as parasite and host. or Among plants by far the greatest number of parasites. which are living upon it. the host is said to FIG. as applied to both plants and animals. the dependent upon some external source of food. FUNDAMENTALS OF BIOLOGY do not. 223. as of saprophytes. however. all found among the bacteria and of which lack chlorophyll and are consequently Parasitic bacteria all the other fungi that are parasitic confine their attacks to plants. X 100. It should be understood that the disease is merely a response on the part of an organism to the presence of a parasite A disease manifests itself by symptoms. as cases of parasitism. but nearly . If injured sufficiently to cause some marked disturbance in its normal functions. has a rather limited meaning. Cross section of a radish leaf infected with a parasitic fungus (Albugo Candida). The mycelium forces its way between the mesophyll cells arid produces spores beneath the epidermis.322 plants. be diseased. Parasitic Plants. abnormalities in function or structure caused both. live on both plants and animals. are other fungi. consider the relation between a predatory animal and its victim. which finally ruptures and liberates the spores into the air. Among the many plant diseases caused by fungi other than bacteria. thus forming a blister on the surface of the leaf.

224. corn smut. potato rot of stone fruits. 323 may be mentioned: downy mildew of grape. diphtheria. tissue. AND SYMBIOSIS following blight. cabbage rot. 224). tuberculosis. cucurbit wilt (attacking melons.SAPROPHYTISM. resulting in the formation of tumors or galls (Fig. cholcria. Of the bacterial diseases of plants' the following are widespread: pear blight. 26 and 223). and crown gall. of man and of the higher animals are caused by bacteria. and tetanus (lockjaw). and wheat rust. lives within the tissues of the host but in most parasitic fungi the mycelium and merely produces spores FIG. The gall consists of on cherry twigs by the "black-knot" mycelium and hypertrophicd host Some parasitic fungi cause at the surface (Figs. Some fungi are external parasites. Most of the diseases squashes. Galls produced (Plowrightia morbosa). the following being well-known examples: typhoid fever. PARASITISM. the mycelium living on the surface of the host and sending short branches (haustoria) into brown the epidermal cells. pneumonia. . a hypertrophy of the host tissue. and cucumbers).

etc. Animal parasites different groups. worms of various Some are parasitic only during part of their life. leeches. a sood-plant parasite. Mistletoes live on various kinds of trees. The Parasitic o d d o r (Cnscuta). Although some kinds of mistletoes 226). flat worms. lice. such as bedbugs. The very plants (Fig. all but. arachnids. such as roundinsects. Hurker-liko processes are sent into the host. them nourishment is animal parasites fleas. the host only ticks. absorbed. the parasite has no with the soil and. a relative of the morning-glory. I) Animals. twining about a poplar stem. and vertebrates. crustaceans.324 FUNDAMENTALS OF BIOLOGY Only a few parasites are found among the higher plants. Except in connection the seedling stage. yellow orange. 225). Short. but Some are very rare among the last two. attacks toes. etc. natural 225. and the are able The latter have green leaves to make at least part of their own food soil. are found among many protozoans. annelids. others are permanently parasitic and cannot exist apart from . others are only partially so. they send suckers into the vascular tissues of the host (Fig. worms. asites. are entirely parasitic. two conspicuous examples being the dodder (Cuscuta) and the mistleThe dodder. such as is made with The most highly when the many specialized parasites are protozoan parasites.. having no connection with mistletoes obtain their entire salts supply of water and mineral host. Like the dodder. may be external. mollusks. lacking chlorophyll almost completely. mosquitoes. is unable to carry on photosynthesis. and through size. practically the host and sends short sucker-like processes into it. and many of these are partial or occasional par- since a contact parasite is feeding. internal ones. from the seeds of mistletoes are sticky distributed arc and by becoming attached to the feet of birds. or leafless stem twines about slender. Fm. These various woody and herbaceous seed penetrate the vascular system. absorbing from it both food and water. kinds.

. Such degeneration has come about as a result of the protecetc. food-getting. Correlated with thoir easy life. having lost their organs of special sense. 325 Some can live on a variety of hosts. many animal parasites. partic- ularly internal ones. one-third natural size. (below) a closer view of another species (P.SAPROPHYTISM. longispicum) growing on a poplar stem. A mistletoe (Phoradendron mllosum) growing on an oak tree and FIG. are sluggish and structurally degenerate. AND SYMBIOSIS their host. 226. others only on one kind of host or on a succession of several specific hosts. PARASITISM. locomotion.

If a malarial patient is bitten by an Anopheles mosquito. One of the greatest discoveries of modern medicine is that organisms which cause certain human diseases are carried by insects. When the infected insect bites another human being. the organism that invades and feeds pointed size. stages some parasites are free living and are often structurally more complex than when mature. The most highly specialized parasites require two different hosts to complete their life cycle. as the two examples given below illustrate. as every 48 or 72 hours. There they undergo a series of finally resulting in the production of a great spores of a different kind. toms occur at regular intervals. As it of a diseased female mosquito of the genus Anopheles. which goes through a rather complicated life The only mode of infection is through the bite cycle (Fig. cell consists of a minute. Malarial Parasite. the organism gradually increases in coming to resemble a small amoeba. 227). waste products resulting from the destruction of the corpuscle as well as those arising from the parasite itself. The parasite that causes malaria in man is a minute protozoan called Plasmodium. and this fact has been known in connection with the transmission of malarial fever only since 1898. upon one of the red blood corpuscles. evident that the only way in which a man can contract malarial fever is through the bite of a mosquito that itself carries the parasite by having bitten either a malarial patient. it produces a number of small spores that escape. coinciding with the production of spores by the parasite. some of these spores are injected complex changes. As a result. processes of growth and reproduction. slender.326 FUNDAMENTALS OF BIOLOGY tion against enemies afforded by parasitism and of the presence In their larval of an abundant and accessible food supply. many into the blood stream. a number of with the blood which malarial organisms enter the body of the insect it sucks. It is the liberation of this poisonous material that causes the chill and fever charIn some forms of malaria these sympacteristic of the disease. It has long been realized that the . After destroying the contents of the corpuscle. It is and the individual contracts the disease. or a person who has recovered from the disease but still carries some of the parasites in his blood. enters the human body. each entering another corpuscle and repeating the When the spores escape. These represent poisons are liberated into the blood stream.

Ltd." A. During the sexual phase of the life history. in Lankester'* Treatise on Zoology. Diagram illustrating the life history of the malarial parasite (Plasmodium The stages above the Tine occur in the human body. stomach and undergoes changes that result in the formation of a large number of and are then ready Suito's ender pointed cells (XVIII) that make their way to the salivary glands " to be injected into a human being. by permission. and producing spores. & C. .) of the mosquito. Black.. Those below in the body I to V and 6 to 10 show parasite entering human red blood corpuscle. sperms and eggs This encysts in the wall of the mosare differentiated and unite to form a zygote (XII). PARASITISM. malariae). AND SYMBIOSIS 327 u FIG. 227.SAPROPUYTISM. (From Minchin. growing.

is composed of a minute headlike organ (scolex) and a number of flat posterior that become segments (proglottids) larger and larger throughout its length The head is attached to (Fig. The segments pass out of the human body through the digesEach contains thousands of small embryos. t 228. The pork tapeworm (Taenia solium) feet is smaller. but a colony. . the intestinal wall and gives rise to the infect 1 posterior segments by a process of budding.ool"General Wieman. Tapeworms. a cow. tive tract. the of tapeworm. apeworm s c o 1 e x (s) enlarged. Many of them require two hosts in order to complete their life The body of a human tape worm cycle. Tapeworms are par- asites that live as adults in the diges- beifig tive tract of the higher animals. showing adhesive hooks and suckers. o0y" after Leuckart-Nitsche entire "head" or A. parasite bores int(5 the muscles. excretory. rarely exceeding 10 in length. flatworms. as its general appearance would seem to indicate. Some of these embryos may then be swallowed by the alternate host. as the tapeworm is hermaphroditic and self-fertilizing. (From 7. there several species that commonly man. undergoes further development. depending on the species After entering the body of the other host. the of metameres. arid reproductive systems are essentially similar to those of other wall chart. A single specimen of beef tapeworm (Taenia saginatd) may consist of several thousand segments and may reach a length of 20 to 30 feet or more. Thus the "worm" is really not a single animal composed of a number A FIG. which may be a hog.) digestive system. worm. Their nervous. 228). . Tapeworms have no they absorb food through their body wall after it has been digested by the host. ( T he pork Taenia solium) /?.328 FUNDAMENTALS OF BIOLOGY most effective way of controlling malaria is to destroy the mosquitoes that transmit the parasites from one human being to another. or some other animal.

Although a disease is any marked disturbance in the normal functioning of the body. individual is known lifetime of an as acquired immunthe during are be either active or passive. upon the activity of the phagocytes and upon the formation of antibodies. by protozoans. Antibodies are substances in the blood plasma that act in antagonism to specific bacteria or to Their chemical nature is unknown. lysins dissolve microorganisms or cause them to disintegrate. AND SYMBIOSIS and finally 329 becomes encysted. Most infectious diseases of man are caused by recog- nizable bacteria or are caused by analogous organisms (filterable viruses) too small to be seen with the microscope.8APROPHYTISM. They accumulate in large numbers wherever an infection exists. PARASITISM. while opsonins act upon microorganisms in such a way as to render them more easily destroyed by phagocytes. The ability of the body to resist the effects of invading germs or their toxins is known as immunity. Disease-producing microorganisms invade the body chiefly through wounds and by way of the respiratory and alimentary The extent to which the body is harmed depends largely tracts. Immunity may be either natural or acquired. beef. upon their recognized. an infectious disease is one brought about by the entrance of specific microorganisms and their subsequent multiplication within the for the presence of tissues. like malaria. such inherited Resistance developed resistance constituting natural immunity. action. while passive immunity results from the introduction into the body of antibodies produced animal. Active immunity is acquired through the production of antibodies by the individual ity. Some persons more resistant than others to a given disease. In this form it enters the body infected meat is eaten. several different kinds of antibodies are Thus antitoxins neutralize toxins and render them harmless. It may who becomes immune. by another . regardless of its cause. Phagocytes are white blood corpuscles that destroy bacteria by engulfing them. agglutinins cause them to stick together. The eating of raw or insufficiently cooked pork. Depending their toxins. In certain diseases the toxins arising stimulate the its body to produce from the causal organisms more antibodies and thus increase resistance to them. Immunity. Some. or fish is generally responsible of man when tapeworms in the human body.

does not confer Active immunity to certain diseases is not dependent and subsequent recovery but can be induced by upon the introduction into the body of preparations containing dead immunity. and yellow fever. epidemic meningitis. results in an extremely mild infection that confers an immunity usually lasting from 7 to 10 years. In addition to typhoid fever. called antigens. cholera. measles. cholera. infection germs. Recovery from such as tuberculosis and malaria. typhus. the amount is often insufficient to effect a cure unless serum. lasting from 5 to 8 years. This is accomplished by rubbing into a scratch made in the skin a preparation containing the living germs (virus) of smallpox after they have been weakened by growth in the body of a calf. supplemented by antitoxin from an outside source. plague. immunity may be acquired by artificial means in tetanus (lockjaw). smallpox. Such preparations. Passive immunity to diphtheria can be secured of diphtheria antitoxin contained in the blood by the injection serum of a horse that has developed active immunity by repeated doses of diphThis antitoxin gives only temporary immunity and is used chiefly as an aid . result in the formation of antibodies. plague. The animal contracts the disease ("eowpox") which. For recovery after the disease has theria toxin. mumps. chicken pox. recovery from some ing immunity from a second attack. fever. Active immunity to smallpox can be induced by vaccination. SYMBIOSIS Symbiosis is an association in which two different kinds of organisms live together to the advantage of one.330 FUNDAMENTALS OF BIOLOGY diseases produces a lastSuch diseases include As is well known. weakened germs. when transmitted by vaccination to a human being. rabies (hydrophobia). measles. can be secured by injecting into the body a suspension of dead typhoid bacteria. already been contracted. a preparation containing the products of growth of the diphtheria germ so modified as not to produce any toxic effects. or oven for life. smallpox. active immunity to typhoid fever. as in horse Active immunity against diphtheria is now most com- monly obtained by the administration of diphtheria toxoid. scarlet fever. and probably also in whooping cough. typhoid other diseases. and diphtheria. scarlet fever. Although the toxin formed by the diphtheria germs stimulates the body to produce its own antitoxin. or weakened toxins. but in such a .

but here we be interested merely in a few well-known examples chosen to illustrate the general situation. but some (Fig. Sometimes there is mutual benefit without injury to either. a ciustose foim (Placodium) growing on rock. This is clearly indicated by a cross section through the lichen body (Fig. PARASITISM. a lichen (Cladonid) that grows erect upon the ground. are more conspicuous. 229). D. tree trunks. Lichens. or if injured some compensating benefit from the association. C. B. number shall of kinds of symbiosis are often recognized. but an alga and a fungus living together in symbiotic relationship. The greater part of the lichen is composed of a dense mass of tangled fungous . as its appearance suggest x. 230). a foliose type (Parmelia) growing on bark. 229. A. dead wood. a branching form (Usnea) that hangs from the limbs of trees. and on the ground Their usual color is gray or grayish grqen.SAPROPHYTISM. Lichens are small plants of various form commonly seen growing on rocks. natural size. AND SYMBIOSIS 331 it way that the other is not materially injured. A lichen is not a single plant. Group of common lichens. and so a receives FIG. There are many different degrees of mutual dependence.

X 500. however. formed Nitrogen-fixing Bacteria. As has been stated. while the fungus gives moisture to the alga. Cross section of a lichen body (Physcia). 230. which the alga could not live. irregularly or in are the cells of the alga either scattered The fungus lives upon the kill it. FUNDAMENTALS OF BIOLOGY among which a definite layer. Thus the association between the two lichen components is one of mutual advantage. nitrates are in the soil both by the action of the nitrifying bacteria . In fact. At the same time. showing cells of the alga (above) surrounded by a mass of interlacing fungous filaments. the alga is benefited in that the fungous body readily absorbs and holds moisture without alga as a parasite but does not FIG. The alga supplies the fungus with This reciprocal food. the alga seems to be but slightly injured. relation enables lichens to grow in dry.332 filaments. exposed situations where neither the #lga nor the fungus would be able to live alone. merely losing some of the food that it makes.

158#). Root sysof a young hoan fact that. like the saprophytic 231). Aphids. and this is then available for other green plants. some of the nitrogenous material becomes available for within their use by the legume. particularly legumes. peas. alfalfa. tem FIG. forms mentioned above that live free in the arc unique in being able to absorb free nitrogen from the air in the soil and to "fix" it. 231. Although some of the latter arc saprophytic upon organic matter in the soil. 318). which convert free nitrogen to nitrates (see p. are small oval green bugs that sucl* the juices of plants (Fig. or plant lice. They are often seen on young tender shoots." of which ants are very . but some kinds attack roots. Aphids secrete a sweet substance from their bodies called "honeydew. bacteria invade the roots of the legume The and live as parasites in the root tissues. etc. beans. When the legume dies. For obvious reasons the relation between the nitro- growth is nitrogen-fixing bacteria one-half natural size. animals is The relation that exists between these a sort of slavery and is a real case of symbiosis because both members of the association are benefited. when cultivated in sterilized (Phascolus) with numerous tubercles in which live. and by the 333 nitrogen-fixing forms. forming tubercles or nodules (Fig. gen-fixing bacteria and the legume is often called reciprocal parasitism. even when nitrate abundance. the roots undergo local enlargement. a considerable amount of nitrogen is added to the soil. Ants and Aphids. to form nitrogenous compounds soil. that is. After this process has been going on for a while. the cultivation of legumes is an important agri- As a means That legumes have come cultural practice. AND SYMBIOSIS upon ammonia. to depend upon nitrogen-fixing bacteria for their supply of nitrogen is shown by the soil. upon carbohydrates As a result of their presence. PARASITISM. others enter into a peculiar relation with certain seed plants. such as clover. of restoring nitrates to impoverished soils. These bacteria. lupines. salts are present in very feeble.SAPROPHYTISM. own cells.

and legs prothe abdomen. The advantage of this association to the anemone is that afford. Some species of hermit crabs place sea anemones on their shell. and and devoid of appendages. which fits inside the shell. their cephalothorax. claws. For this rea-son aphids are often called "ants' cattle. and thus taken into new feeding truding. Some kinds of ants take the aphids to their own nests during the winter and feed them in order to secure a constant supply of honeydew. grounds. defend the aphids against attack and take care of them in other ways. becoming ." Hermit Crabs and Sea Anemones. ants may be seen running fond.. Hermit crabs live inside empty soft snail shells. and thus gain the protection that these creatures. back and forth between the aphids and the ant nests gathering The ants the sweet secretion and carrying it to the larvae.334 FUNDAMENTALS OF BIOLOGY Wherever aphids are found. by reason of their numerous stinging cells. it is carried about by the crab.

by a process of orderly change. It maintains that no changes. and that consequently each kind of organism is fixed and immutable. 335 The latter . stages. has produced convincing evidence as to the reality of evolution the process whereby modern forms of life have been derived from All earlier. views have been held with respect to the past conditions of organic nature. which superseded by conception of a static world. represented by an unbroken succession of individuals. Life. The older one is that the earth and its inhabitants came into existence in a form like that in which they now exist. the study of evolution is not concerned with the origin of life but only with its development. Geology teaches that the earth. or how the first life. has been continuous from the beginning. one species never giving rise to another.ones. and.CHAPTER XX THE FACTS OF EVOLUTION study of organisms. no one knows. appeared. except perhaps minor . arose. all are related by descent. during its formative was lifeless that primal conditions were not favorable existence of organisms. this dominated human has been the thought. to this process of gradual change they are the modified descendants of preexisting species. Its origin is taken for granted. for the When. which has long recognizes the reality of the process of evolution. forms more and more complex. since all organisms have had a common origin. the multitudinous species of plants and animals now inhabiting scientific The the earth owe their present condition of structural organization . in all its many aspects. because of present limitations of our knowledge. dynamic view. where. have occurred since life began. and that from them have come. simpler forms through the operation of natural laws. all But regardless of how the first living forms available evidence indicates that they were extremely simple. In fact. Two Owing to the influence of modern science.

smell. which is itself supported by not a single fact. any case. in general. The hour hand on a clock moves much more slowly than the minute hand. but is actually proved by the historical record of life that has been preserved in the rocks in the of fossils. geology. not only on the earth. as we have soon. so obvious. Although evolution has been mainly progressive. The fact of racial development. although not nearly is just as real. but not necessarily so. The gradual increase* in structural complexity as an organism passes /rom the fertilized egg through successive stages in its growth is self-evident. while racial development. From an extremely simple original condition. called comparatively rapid and of but brief duration. characteristic of civilized man. is The dulling of the senses of sight. is ontogeny. in many cases it has operated in the opposite direction and has led to a reduction or loss of parts. these alone operate in the process of orderly change that characterizes the physical world constitutes what is known as that. Retrogressive changes. so far as and The Recognition of its occurrence is fundainorganic evolution. physics.336 is FUNDAMENTALS OF BIOLOGY significance not only supported by a vast array of facts that arc without on the basis of the former view. form All the physical sciences testify that everywhere in the universe dynamic conditions prevail. which given. However. No one questions the fact of individual development. has been associated with in the development of the brain. living forms have. has proceeded with extreme slowness over an inconceivably vast extent of time. but on all heavenly bodies. Organic evolution is essentially the same process in living things. and hearing. or to other degener- ative changes. followed a course of changes that lias led to everincreasing complexity of structural organization and specialization of bodily functions. is is sciences also demonstrate that the universe ruled The physical by natural production of changes. yet moves just as certainly. Perpetual change is going on everywhere. and chemistry. They are often correlated with progressive changes in another direction. are usually associated with parasitism. Many other examples could be whether evolution has been mainly . known. mental to the sciences of astronomy. or phylogcny. Per- manency laws. Individual development. in nature is purely a relative term.

it should be realized that any reluctance one might have in admitting the occurrence of evolution. of biology. or of any other natural process. etc. government. axiom warfare. Moreover. branches of learning. the only rational explanation of a great mass of facts that whatsoever otherwise would be utterly meaningless. probably has had a more profound influence on human thought than has any other At the present time it dominates all scientific generalization. as will be seen in a subsequent chapter. One does or does not "accept" evolution in the same sense as any other scientific generalization based on demonstrable facts. The doctrine of descent with modifundamental axiom of biology. There is no controversy among biologists that organisms have evolved. evolutionary interpretation of nature scientists of recognized standing is and has boon accepted today by all since the middle It is universally accepted because it offers of the last century. customs. As a scientific prinThe ciple it is as well established as the law of gravitation. an evolution of languages. although. supported scientific evidence. Another misconception is that the knowledge of evolution that one obtains through a study of nature is antagonistic to This is based on a failure to differentiate between the religion. Briefly stated. A great deal of confusion exists in the popular mind concerning evolution. We recognize. fication is a it is 337 always a process of racial Importance of Evolution. although there are many theories relating to how it operates. even those having no relation to science. for example. One of the most prevalent thafc is that evolution faith. is the subject of a creed It one accepts on something should be understood that the principle of evolution. It is not a theory. in no way detracts from its reality. the establishment of the principle of organic evolution has revolutionized man's conception of the universe and of himself. is by an overwhelming mass of a logical explanation of the facts of nature. siderable difference of opinion as to The establishment tal of the principle of evolution as a fundamen- which represents the greatest intellectual accomplishment of the nineteenth century. religious thought. there is conhow they have evolved. . and as a result many false ideas regarding it has become current. Popular Misconceptions.THE FACTS OF EVOLUTION progressive or retrogressive modification.

according to any other interpretation they arc rendered wholly unintelligible. living contemporaneously with its modified descendants. each appeals to different Science does not deny the existshows that the method of creation is merely To say that the world has developed in accordance evolution. of is to develop the consciences. or from some other This misconception is based existing species of ape or monkey. Millikan says: without prejudice or precona the of facts. . common ancestor unlike Thus the relationship between man and the either living form. and the procany kind. The purpose ception of other hand. it also indicates that each has developed independently along a divergent line from some extinct. It is of man from understand how the process of evolution has Except in rare instances one form of life cannot be operated. on the of science is to develop. The dis1 tinguished American physicist. Not only do numerous facts of biology support this explanation. the laws. but none has ever been discovered that refutes it or warrants on a failure to Part of a joint statement issued religious leaders and scientists. Nature of the Evidence. and the aspirations mankind. call it what we may. Nearly all the countless species of plants and animals of the past are utterly extinct. basis than that of descent with modification. knowledge esses of nature. as ancestral to another that is contemporaneous with regarded 2 it. only when one takes upon it aspects of human the functions of ence of a creator. There can he no It is real conflict because each properly occupies a itself different sphere. with natural laws does not deny the existence of an unknown guiding influence. the ideals. 1 and signed by a group of 31 leading 2 Only a relatively few cases are known where an ancient type of organism has persisted in a comparatively unchanged condition into the present. but strictly collateral. thought. the other that difficulties arise. The principle of organic evolution is an explanation of demonstrable facts drawn from all fields of These facts cannot be explained on any other biological science. modern apes is not lineal. the chimpanzee. generalized. Robert A.338 FUNDAMENTALS OF BIOLOGY functions of science and the functions of religion. popularly supposed that evolution implies direct descent the gorilla. Although all the existing evidence indicates that both man and the apes have evolved from a lower type of life. The even more important task of religion.

Thus the dog family includes the dog genus (Canis) and the fox genus (Vulpcs). and although never exactly alike. It should be noted that the scientific name of an organism is the name of its The position of the genus combined with the name of its species. show less variation among themselves than they do with members of any other similar group. Classification. for example. ) r - . genus includes one or more and in some species still smaller divisions. the gray or timber wolf of North America (Canis occidentalis) the prairie . A phylum may be divided into several subphyla..THE FACTS OF EVOLUTION any alternative explanation. The carnivores. (the chordates) Phylum Chordata Class Subphylura Vertebrata (the vertebrates) Mammalia (the mammals) Order Carnivora (the carnivores) Family Canidae (the dog family) Genus Gfonis . It includes individuals that closely resemble one another in form and structure.. An order is composed of still smaller groups called families. the jackal (Canis aureus). and mammals. The major groups are called phyla. each constitute an order mammals. 232). The species is the unit of classification (Fig. is an assemblage of related genera. include the dog family. The generalization and is evidence upon which the principle rests comprises "the facts of evolution/' Some of the more striking classes of these facts we are now ready to consider. etc. bats. reptiles. Each class consists of smaller groups called orders. birds. in turn. > Species famihans ) The common dog . the bear family. We have already seen that all members of the plant and animal kingdoms fall naturally into a number of distinct groups that can be arranged in an ascending series. A family. Thus the vertebrates are a division of the phylum Chordata and consist of five main classes: fishes. etc. dog in the animal kingdom is indicated below. primates. . are recognized. but more commonly directly into classes. wolf or coyote (Canis latrans). The carnivores. . etc. called varieties. ungulates. The dog genus includes not only the various breeds of the common domestic dog (Canis familiaris) but the European wolf {Canis lupus).. Finally each species. the more important of which we have already considered in both the plant and animal kingdoms. of rodents. evolution stands as 339 of Thus ~the doctrine scientific organic a great accepted today as such. the cat family. amphibians.

The basis of classification. reel oak (Q. bur oak (Q. by FIG. macrocarpa) . blackjack oak (Q. black oak (Q.) . B. A. rubra). white oak (Q. virginiana). D. E. "Practical Botany. after Hough. live oak (Q.340 FUNDAMENTALS OF BIOLOGY group of leaves and acorns illusoak (Qucrcus). (From vdutina)." Ginn and Company. marylandica) . F. A trating some of the differences between six species of permission. alba). 232. Bergen and Caldwdl. C.

some higher. The members of every natural group of organisms have common structural features because they have had a common origin. a dog is more like a wolf than it is like a This clearly indicates cat. on the contrary. ture extends throughout all parts of the mammalian body. have the same relation to one another as do the branches on a tree. the closer is their relationship. and all can be traced to a common trunk. For example. What is the significance of fundamental structural resemfact. some lower on the tree. then. the "higher" groups a vastly greater amount. it has previously been pointed out that the limbs of mammals show the same fundamental plan of structure. the differences among them having arisen through each having subsequently followed a divergent course of evolution. And relationIt is no mere ship denotes descent from a common ancestry. coincidence that a tiger and a leopard are strikingly similar. arc not like rungs on a ladder. and that both the plant and animal kingdom . 308-312). a wolf and a fox. blance? It means but one thing relationship. and that the differences seen among life the various members of the group are related to Basic similarity in struchabits (see pp. but more like a cat than a horse. mere fact. each representing a definite amount of progress in a single direction but. In diverse a comparative study of the members of any group of organisms shows that. dog and the wolf must have been less remote in time than the one common to the dog and cat. that organisms can be classified on the basis of structural similarity. Thus the greater the degree of basic resemblance between any two organisms. despite many superficial differences. in each case. Plant and animal groups. Existing groups are at the ends of twigs. however. their bodies are built according to the same general plan.THE FACTS OF EVOLUTION What is 341 It is the basis of this scheme of classification? fundamental structural resemblance. The branches represent divergent The lines of descent. Their resemblances. The "lower" groups of plants and animals are merely those that the ancestor common to the which have undergone relatively little facial modification. As every one knows. and consequently the less remote the common ancestor from which both have sprung. or a duck and a goose. while the ancestor common to the dog and horse (and to other mammals as well) must have been still more distant. are due to actual organic relationship.

1 B FIG. In fact. Most plants and animals possess certain Vestigial Structures. 234. Flowers of garden asparagus. after H. having a vestigial shell covered by the mantle (w). or less degenerate condition. a snail (Helix).342 FUNDAMENTALS OF BIOLOGY can be represented in the form of a tree. natural size. 233. parts that perform no useful function and are present in a more FIG. with fully developed (From Bergen and CaldweJl. The flowers of the common asparagus are of two kinds: one having perfect stamens and a rudimentary pistil. staminate flower.) . pistillate flower. it has no other rational explanation. tures. with perfect stamens () and rudimentary pistil (rp) #. . Botany. These are called vestigial struc- Their presence can be adequately explained only by assuming that they were more fully developed and functional in the organism's ancestors. the other with . natural size. B. A. a mollusk with a well-developed shell.' Ginn and Company. by permission. A few examples will be given. Mueller. "Practical pistil (p) and rudimentary stamens (rs). signifies that evolution has taken place. a slug ( Ariolimax) a closely related form. A.

The snakes and lizards belong to the same group of reptiles. The only reasonable interpretation that can be placed on this fact is that the ancestors of the asparagus had flowers in which both the stamens and the pistil were functional. less The presence testifies of this use- shell that the slug and hind limbs of the python. limbs are absent. is indicated by the presence of vestigial hind limbs in certain existing species of snakes (Fig. The power of moving the ears so as to catch . In many of the lower vertebrates there is present a third eyelid (the nictitating membrane) consisting of a thin translucent membrane that may be drawn diagonally across the eyeball. however. The asparagus also has rudimentary leaves in the form of small scales that are entirely functionless. in fact it is really a snail with a shell. of a thin flat plate embedded in the mantle. Obviously an adaptation to compensate for the reduced condition of the leaves. X y. man . 235. by permission. "Darwin and after Darwin." Open Court Publishing Company.THE FACTS Q EVOLUTION 343 rudimentary stamens and a perfect pistil (Fig. but correlated with the special mode of locomotion assumed by the snakes. is for photosynthe- the fact that the stems are green and highly branched. (From Romanes. the branches being very fine. Vestigial pelvic girdlo the animal's back. 236). It is part of their plan of organization to have four limbs. That they have been derived from limbed ancestors. 233). In this structure is represented by a remnant called the semilunar fold (or plica semilunaris) situated at the inner corner of the eye (Fig. The shell is lined with a thin muscular sac called the In the slug the mantle mantle. and to provide sis. 235). The garden slug is a mollusk closely related to the snails. consists of a small oval patch on FKJ.} has been derived from snail-like ancestors. the shell. 234). reduced Snails greatly possess a spiral shell into which the soft body can bo withdrawn (Fig. as they have many structural features in common.

236. membrane (N) in the owl and horse compared with (From Romanes.) Gorilla FIG. "Darwin and after Darwin. (From Romanes. its The nictitating rudiment in man. Open Court Publishing Company.) of gorilla Sacrum . 37. showing the rudimentary tail bones of each. by permission.344 FUNDAMENTALS OF BIOLOGY Owl Horse PLICA SEMI LUNAR is Man FIG. Man compared with that of man. by permission." Open Court Publishing Company. "Darwin and after Darwin".

There are present. while all of the higher groups have both features. every its Thus it is human being a self-evident fact that. that this cell represents an individual in the simplest possible condition of structural organization. the sequence of its embryonic stages is indicative of the development through which an animal has gone in the course of its evolution. Or. are pergastrula stage. but that it was once present is evidenced by the occurrence of feebly developed ear muscles lying close against the head. representing a rudimentary tail that is well developed during a portion of early embryonic life (Fig. In man this power has become lost. the parallelism between individual and racial development is perfect. for it is always only approximate. like other is unicellular at the beginning of existence.THE FACTS OF EVOLUTION 345 sound more effectively is well developed among such mammals dog and horse. 233-236). others in the of cleavage The The coelenterates. manent gastrulas. They all testify to the reality of evolution. at the lower end of the spinal column. having neither mesoderm nor a coolom. The members of the next group. The recapitulation principle is well illustrated among insects. Attention has been called repeatedly to the fact that every organism which reproduces by the sexual method It is apparent begins its existence as a single cell the zygotc. as often stated. but multicellular organisms only temporarily. onto- geny (the development of the individual) recapitulates phylogeny This is a statement of the (the development of the race). It should not be taken to mean that principle of* recapitulation. The examples cited above of vestigial structures are only a few selected from thousands that have been described. We have seen that the annelids and arthropods have many struc- . over a hundred from man alone. In a general way. series of early embryonic stages involved in the processes and gastrulation are essentially alike in practically We have seen that all multicellular animals (see pp. it will be recalled. some of the lower animals proceed only a short way in their development. metazoans. have mesoderm but no coclom. brief Embryonic development is a and condensed repetition of a series of ancestral stages through which the race has passed. Embryology. a condition in which unicellular organisms remain permanently. some remaining in the blastula stage. the flatworms. a short row of as the coccygeal bones. 237).

This figure also shows that members of different classes begin to ment. but these are always where embryos representing each vertebrates are shown. worm-like ancestors. in Fig. of the five great classes of manifest conspicuous differences much earlier in development . 238. as development proceeds. (After Romanes. bee. or beetle is distinctly worm-like. "Darwin and after Darwin. fly. by permission. relationship persist during later developFor example. not only in general appearance but in a number of more fundamental respects (Fig. differences become more pronounced. This strongly indicates that the insects have evolved from 165). indicating descent from a common ancestry. 238. A series of vertebrate embryos at three Although the the earliest stages in embryonic development are it is essentially similar in all metazoans. resemble one another more than they do any of the other vertebrates. Note that the four mammalian embryos. but. striking. at each stage. The larval stage of a butterfly. Fish Salamander Tortoise Chick Calf Rabbit Man comparable and progressive At first all are much alike. The life history of any member of one of the higher groups of insects substantiates this interpretation.346 tural features in FUNDAMENTALS OF BIOLOGY common and thus are closely related.) FIG. stages of development. only among members of same group that resemblances is apparent by their general structural similarity." Open Court Publishing Company.

347 denoting descent from an older common stock. that. Similarly the human brain. Intergrading Species. But. Likewise. when gill slits are present in the human embryo. intergrading species. or become modified to form other structures. hawthorns. The latter become functional only in the fishes and amphibians. as has been seen." Typical members of one species be very different from typical members of another. some genera difficult to determine their limits. roses. Before evolution had become an established principle of biology. in common with all vertebrates. as in birds its and in other mammals. some show such a great range of variation that it is very In other words. fishes. on the basis of the principle of evolution. For example. 238). persist as vestiges. Moreover. and mammals. in embryonic development. it would account for the occurrence of intermediate forms. but their appearance in all vertebrates is indicative of descent from aquatic ancestors. In the animal kingdom. passes through a series of stages corresponding to adult conditions in the lower vertebrate groups.THE FACTS OF EVOLUTION than members of the same class. species. interoaks. and taxonomists differ among themselves as to where the lines of demarcation are to be drawn. It was soon found. the human embryo has a tail as well developed as that of any of the other vertebrates (Fig. two chambered The heart then passes through a three-chambered stage. it has gill slits. Embryology teaches that many structures which are permanent in the lower members of a group appear only in embryonic stages in the case of the higher members. In all such cases trie limits of each species are more or less arbitrarily defined. If all species difficult to be were fixed and incapable of modification. and then either later disappear completely. individuals that show some of the characters of both This is true of the asters. the heart is and the circulatory system distinctly fish-like. grading species are especially common among insects. during an early period of prenatal development. and finally becomes four chambered. it was supposed that every species of plant and animal has always existed in its present state. although many species seem to be distinct. characteristic of amphibians and reptiles. however. and that each one is entirely distinct from all others. and many other plants. but often individuals that are intermediate between the two may be found. genera exhibiting . exhibit " may birds. willows.

by permission. the probable ancestor of the group.) of Botany" P. Brussels sprouts. cabbage (Brassica kohlrabi. C. broccoli. wild cliffoleracea). 239. E. kale. Blakiston's Son Horticultural plants that have arisen from the wild cabbage. A. cauliflower. Company.Fia. D. B. F. G. "Fundamentals & 34S . (From Gayer. cabbage.

dogs. and from this center of The subsequent dispersal spread as far as conditions permitted. but also cauliflower. broccoli. kale. example. establishment of barriers created. in that their origin is lost in obscurity. fication. In many instances such profound changes have taken place that it is impossible even to recognize the wild ancestral form. it is difficult to realize that each group has evolved from one or If species were incapable of modisuch striking changes as these would not have been possible in such a comparatively short time. etc. From this single species man has developed not only all the different kinds of cabbages. In general. Many have been under man's influence since prehistoric times.THE FACTS OF EVOLUTION 349 intergrading species may be merely those in an active condition of evolution at the present time. parts of Australia. Cultivation and Domestication. bution. being kept from by a barrier of some sort. for so long a period. kohlrabi. it has been found that similar but distinct species often occupy adjacent parts of the same region. poultry. or came in from elsewhere. This fact can be satisfactorily explained by assuming that the different species have sprung from a common ancestor that originated in a particular part intermingling of the region. etc. such as several original wild species. Everyone is in agreement that all cultivated plants and domesticated animals are the modified descendants of wild species. Geographical Distribution. . fact. the wild cabbage still grows in Europe but bears little superficial resemblance to its cultivated descendants (Fig. a desert. such as a mountain range. 239). a body of water. striking facts are often revealed.. different kinds of pigeons. The facts of geographical distribution present additional evidence in support of the principle of Some plants and animals are cosmopolitan in distrievolution. local groups of * individuals that evolved along divergent lines and finally became specifically distinct. but most species are restricted in their natural range to That peculiarities in distribution cannot well-defined regions. horses.^South America. cattle. however. be explained on the basis of climatic factors alone is shown by the frequent occurrence of vastly different types of plants and animals in regions widely separated but climatically similar. When one considers all the Brussels sprouts. Where such recogni- For tion is possible. by isolation. and Africa. etc.

that some groups are almost never present. a volcanic group situated about 600 miles west of Ecuador. Regions recently separated. The endemic species. native mammalian fauna of Australia consists almost exclusively of monotremes and marsupials. The discontinuous distribution of the same group in widely separated regions can be explained on the basis that their distrifossil record. These ancient forms survived in Australia. and so must have been derived from ancestors accidentally brought to the islands from the mainland at rare intervals by wind or waves. insects. . tapirs live only in southern Asia and in Central and South America. although distinct. to the length of time the barrier has been in existence. such as the British Isles and Europe. This can often be proved by the For example. show a great general resemblance to South American forms. but nearly all the species of land birds. but during relatively recent geologic times they ranged over eastern Asia.e.. groups that in geologic times were widely spread over other continents. Thus the Special cases of distribution are often interesting.350 FUNDAMENTALS OF BIOLOGY Substantiating this interpretation is the additional fact that the degree of dissimilarity existing between related species separated by a barrier is directly proportional to its geologic age. whereas regions long isolated. in common. bution tvas once continuous. tains no amphibians and no mammals except bats. like Madagascar and Africa. . The life of oceanic islands is peculiar in that fewer kinds of organisms are present than on continental areas of similar size. because there they were kept out of competition with higher types that evolved elsewhere. The Galapagos Islands. have few forms of life i. has sea birds of the same species as are found on the mainland. show great similarity in their fauna and flora. Later they became extinct except in the two widely separated regions where we now find them. and that a great number of endemic species occur species found nowhere else. and Europe as well. North America. The native fauna conreptiles. which has been isolated since the close of the Mesozoic era. and snails are endemic.

reality of organic evolution in a very striking and direct way. Actual Organic Remains. the carcasses of mammoths have been found frozen in the soil of the arctic In some cases the flesh was so well preserved that it tundras. Nature and Formation of Fossils. according to their method of formation. Fossils are the remains of plants and animals that once lived upon the earth. higher and higher types of life came into being from simpler earlier types by the natural process of evolution. for the soft parts nearly always decay or are eaten by animals. special These are simply dead organisms or their parts that have been preserved from decomposition means and consequently have undergone For change. fossils are types. a kind of fossil resin. and that is only a relatively few of the countless number of individuals left that have lived on the earth have ence. are a direct record of evolutionary changes. The preservation of why organic remains occurs under exceptional conditions. in Siberia and Alaska. written in the rocks. fossil The crust were formed.CHAPTER XXI THE of fossils LIFE OF THE PAST Paleontology deals with the history of life as revealed by a study the remains of organisms of the past found embedded The " testimony of the rocks" demonstrates the in the earth. record proves that a progressive development of plant and animal life has taken place throughout the course of geologic As the successive strata that constitute the earth's history. preserved by the tar into which they fell by some little 351 . example. occur as three different In general. as follows: ' 1. could be fed to dogs. Generally they are the hard parts of organisms. The famous asphalt beds in Los Angeles have yielded the actual bones of many extinct mammals. any trace of their exist- found in sedimentary rocks and. The facts of paleontology. Many beautifully preserved insects have been found in amber.

240. Group of bones of the imperial maman asphalt pit at Rancho La Brca. etc. OF BIOLOGY fossils for Actual organic remains arc the most favorable study but unfortunately are very rare. It is chiefly hard parts that have been preserved in this way. 242 and 243). When the organic matter of a dead plant or animal decays but is replaced. (Courtesy of LOB Angeles Museum of History. (Fig. particle by particle. Petrified fossils show not only external form but often internal structure as well. Petrifactions. Los Angeles.352 (Fig. shells. such as wood. If These are the most common kind the body of a dead orjrimNm falls . a petrifaction is formed. California. 3. 241). Natural Molds and Casts. FUNDAMENTALS 240). by mineral matter dissolved in ground water. FIG. of fossils (Figs. bones. teeth. In such cases thin sections of them can be prepared for microscopic study. Science.) ^ ^ 2. moth in Actual organic remains. and Art.

and other external features of information concerning their internal structure. at a rate of 1 foot in reliable estimates (F. the impression may be preserved after all the organic matter has been destroyed.. . Petrified logs.but give no shape. near Holbrook. in the Petrified Forest National Monument. chiefly down of highlands by running water. apparent that molds and casts tell a great deal about the size. Division of Geologic Time. W. 241. it will be necessary for the reader to understand how the relative age is determined of the of life as revealed various layers of rock that form the earth's crust. FIG. Geology teaches that the surface of the earth is constantly undergoing forces transformation as the result of the action of various natural upon it. !^-r !:. Arizona. The by A\< aih<Tin. This cavity may It is later become filled with mineral matter to form a cast. and its suksrquont transi disintegration of rock portation. at the present time. a natural mold results. 1924). of Triassic age. One of the most important sets of changes are those involved in the processes of erosion and deposition. Clarke. the continents as a whole are being degraded. Before considering the history by the fossil record.THE LIFE OF THE PAST into soft 353 mud or sand. and a filling in of result in a gradual wearing According to depressions. If the sediment then hardens to form rock.

carried to lower levels. FIG. Later these sediments may become solidified to form such kinds of rock as sandstone. processes of erosion and deposition that are going on all parts of the world have always been in operation. FIG. 242. gravel. clay. vast portions of what are now dry land have been covered by large bodies of water (Fig. slightly reduced. An impression in rock of a fern leaf (Pecopteris miltoni) from the Upper Carboniferous of Illinois. natural size. 243. 244). etc. Impression in rock of a Miocene fish. sedimentary rocks have been formed. During these periods of subsidence. is deposited as sand. Subsequently these areas have been lifted into the air and once . The eroded material.600 years.354 FUNDAMENTALS OF BIOLOGY 8. As a result of continued degradation and slow sinking of the The today in earth's surface. and conglomerate. shale.

Because sediments accumulate very gradually and vertical movements of the earth's surface take place with extreme slowness. each era into a number of period*. and the characteristic life of each. The accompanying 245) gives the names of the geologic eras. may be divided into epochs. . during which there occurred the earliest known great invasion of land by the oceans." Charles Scribner's udying and comparing the series of stratified rocks in all parts of the world and the succession of fossils that they contain. evident that the various strata of sedimentary rocks forming a large part of the earth's crust have been laid down in It is FIG. while each period again table (Fig.THE LIFE OF THE PAST 355 more exposed to the various agents of erosion. their estimated duration. "Origin and Evolution of Life. 244. The lands were probably all low and the climate warm.) folding. Geologic time is divided into five great eras. and oceans (black) in Upper Cambrian time. except where disturbed by faulting or younger rocks lie above the older ones. the chronological order and. geologists have been able to reconstruct the history of the earth and its inhabitants in a very definite way. enormous stretches of time have elapsed between and during successive periods of deposition. continental seas (gray). (From Osborn. Theoretic restoration of the North American continent (white). by permission. By .

of geologic eras. the oldest rocks were formed satisfactory means devised yet of obtaining . give about total years the time since formation elapsed of the oldest rocks found in the 100 million as the earth's crust. Table More on The recent calculations. they simply stagger the imagination. based present rate of erosion. Reeds. In fact. According to these calculations. 1931) are based on the rate of alteration active elements. have resulted in a new time accepted scale that is widely by modern geologists and paleontologists as the most an accurate estimate of the total length of geologic time and of its divisions. this figure is now thought FIG. the rate at which rivers bring salt to the ocean. 245. length a human period lifetime. The geologist has considerable data enabling him to estimate the relative length of the divisions of but to assign to each an absolute length is a diffiThis arises from the cult matter.356 FUNDAMENTALS OF BIOLOGY Length of Geologic Time. at first glance they seem unbelievable. The geologist deals with time units comparable in their magnitude to the space units employed by the astronomer in measuring the distance between the earth and other heavenly bodies. of integration of the radioactive ele- the radio- ments contained in rocks of known relative age. Because of certain errors in unavoidable the older methods. as compared to of the. geologic time. based the invariable rate of dis- These estimates (revised by C. fact that there of is no certain way age of the determining the earth. Because these time units arc so stupendous.000 years). or even to the of recorded human history (about 5. to be too low. figures in the time scale represent millions of years. The on the or on older methods. A.

and to have remained for a long time in a simple form. were formed. and no one knows anything about their nature. There is no doubt that the duration of the Archeozoic was enormous 800 million years according to reliable estimates. some Archean formations consist of metamoron the earth. comprising the Archean system. and the elapsed time by each of the five eras is as follows: MILLION YEARS Cenozoic Mesozoic Paleozoic 60 135 Proterozoic Archeozoic 355 650 800 2. no certain knowledge is available concerning the early stages in the development of life some evidence. Presumably they were extremely simple. or under what circumstances the first organisms appeared. but beyond that. Many geologists think that it was longer than all subsequent time. and appear at the surface only where uplifted and exposed by erosion. for no one knows when. geologic history when the oldest known rocks. and that by the end of the Proterozoic considerable evolutionary progress had been made. beginning with the oldest. all these ancient rocks. subsequent have undergone intense alteration through vulcanism and great deformative earth movements.THE LIFE OF THE PAST 357 represented approximately two billion years ago. In fact. where. of the existence of life during the Archeozoic. Because life is thought to have arisen during some part of this vast era.000 will Total each of the great geologic eras considered. Although mainly igneous in origin. perhaps more so than any existing organisms." Striking evidence that life existed in abundance at this time is seen in the vast quantities of carbon that occur throughout the the Archeozoic . Moreover. There is phosed sediments. mostly indirect. They constitute a comple#|eries of formations underlying all others. life of The now be briefly THE ARCHEOZOIC AND PROTEROZOIC ERAS The beginning of life is veiled in mystery. very little is known. is designated as the "Age of Unicellular Life. The Archeozoic is the portion of Life of the Archeozoic. to their formation.

fossils . In contrast to the predominating Life of the Proterozoic. and so it was almost as extensive as the Archeozoic. and worm burrows. may which Canada. and because of the subsequent alteration of the Archean rocks by great heat and tremendous pressure. their early evolution during the Proterozoic. Animal undergone from Proterozoic strata. enduring high temperatures that would be Thus these algae may have lived on the earth during the Archeozoic. so far as can be judged. bespeaking an enormously long period of time involved in their formation. Furthermore. however. but are closely related to bacteria. Many of them live in the waters of hot fatal to other organisms. Being probably of a simple and perishable nature. however. Both systems are of immense thickness. The occurrence activity. whatever direct fossil evidence they might once have contained has been largely obliterated. the earliest organisms were not amenable to preservation.358 FUNDAMENTALS OF BIOLOGY Archean system in the form of graphite. It is of limestone is also generally indicative of organic thought that the Grenville limestones of eastern are many thousands of feet in thickness. have been formed by the precipitation of lime from sea water through bacterial action. Figure 245 shows the estimated duration of the Proterozoic to have been 650 million years. the simplest of all of life. the occurrence of what seem to be fossil blue-green algae has been reported from Archean rocks of northern Michigan. comprise about the only remains that have been found. are both scarce and fragmentary a few protozoan shells. The Proterozoic has been termed the "Age of Primitive " because the overlying Cambrian rocks contain Invertebrates abundant remains of higher invertebrates that must have . Recently. Most of the animals of the time were probably soft bodied and hence not suitable for fossilization. It seems certain that this carbo'n must have been derived from the bodies of organisms. the rocks of the Proterozoic are chiefly sedimentary in origin and some of them have under- gone extensive metamorphism. igneous rocks of the Archeozoic. springs and geysers. This discovery is of great interest because the blue-green algae are not only the simplest chlorophyll-bearing plants existing known forms today. some sponge spicules. especially if its surface was hotter then than it is now.

some can directly from water and carbon dioxide (or carbonates) without the aid of chlorophyll and light. In fact.THE LIFE OF THE PAST whatever fossil 359 records might have been preserved have been is largely destroyed by subsequent metamorphism. these plants obtain energy from the oxidation of various inorganic compounds (see p. Extensive sedimentary iron-ore and deposits of Proterozoic age occur in the Lake Superior district. Perhaps they were derived from free-swimming green organisms similar to some of the existing flagellates (see p. since they most closely resemble must algae Simple unicellular animals probably evolved at an early period. many types have been described from Proterozoic strata. (3) Although most bacteria so could not exist in the absence or and are saprophytic parasitic. etc. 318). iron bacteria. sulphur bacteria. Simple blue-green have appeared first. When is the evolution it of the first multicellular organisms occurred unknown. make food of other forms of life. In spite of the paucity of animal fossils. Proterozoic rocks of Montana. 32). with good reason. Instead of utilizing solar energy. but obviously animals must have followed. reaching a great thickness. Early Stages in Evolution. Some of the Proterozoic lime- stones. A later stage in the development of life may have been the appearance synthesis of simple algae green plants able to carry on photo- by the utilization of solar energy. In fact. the appearance of plants. it is thought. The first animals may have bacteria. arisen either during the bacterial or algal stage. while fossil bacteria looking like nitrifying forms have been found in fossil with their formation. a world devoid of bacteria is inconceivable. but . are made up in part of what are considered to be the secretions of calcareous algae similar to those formed of fossil algae by existing blue-green algae. There are three reasons for supposing that bacteria may have been the first forms of life to have existed on the earth: (1) Bacteria are the simplest and smallest of all existing organisms. there abundant evidence of the widespread occurrence of bacteria and algae throughout the Proterozoic. not preceded. (2) Because they are chief agents in effecting the decomposition of all dead organic matter. that bacteria were concerned bacteria closely resembling living iron bacteria have been reported from this region. Examples are the nitrifying bacteria.

as primitive invertebrates were certainly present in the Proterozoic. 246. must have been Fia. During the third great era. But early in the Paleozoic a momentous .) THE PALEOZOIC ERA The little is fossil known record really begins with the Paleozoic. 1931. Table of later geologic time. A. Reeds. since so of the life of the Archeozoic and Proterozoic. much progress was made in the development both of plants and of animals. (Time estimates after C.360 FUNDAMENTALS OF BIOLOGY early. At firsfrall life was aquatic. being probably confined to the margins of shallow epicontinental seas. The figures in the time scale represent millions of years.

since Cambrian fossils are very numerous and diverse. a group of primitive crustaceans. bottom-dwelling forms known as ostraco- . mollusk-like animals of which only a few still survive in trilobites. entirely of marine invertebrates. They are. They were primitive. No trace has ever been found in Cambrian beds of land animals or of verteThe dominant and most highly developed forms were brates. With the beginning of the Paleozoic an extensive development of life had been reached. but greater than the combined duration of the two following eras. but if land plants were present during the Cambrian. only slightly It comprises seven periods that fall into three groups. they have left no fossil record. in most cases. During the Ordovician.THE LIFE OF THE PAST forward step was taken the first land plants and air-breathing animals appeared. each characterized by a definite advance in life (Fig. coeleiiterates. As a result. but algae must have been very abundant to have supported such seems likely that primitive nonsimilar to modern bryophytes. 247). however. and many of ^hese were realized before the era closed. but straight or slightly curved. great possibilities were opened for further evolutionary progress. they were shelled. echinoderms. In also present in the Cambrian seas. total duration of the Paleozoic The years. During the Ordovician. however. They were exceeded numerically. by the brachiopods. It woody plants. was about 355 million more than half that of the Proterozoic. perhaps somewhat display in number of species and individuals (Fig. Fossil algae furnish the only record of the plant life of the period. deeper parts of the ocean. the first fishes appeared. and all of them had a shell like that of the modern chambered nautilus except that it was not coiled. while the largest and most powerful animals of the time were cephaloSome of the latter were 12 to 15 pods. They flourished during the entire Paleozoic and at its close became extinct. nearly all the great invertebrate groups were represented. armored. they made their greatest a luxuriant marine fauna. that is. feet long. Brachiopods were nearly as abundant as trilobites. were becoming established on land. era. Sponges. and true mollusks were fact. a group of true mollusks. At this time the trilobites reached their climax. Cambrian and Ordovician. 246). marine invertebrates remained dominant but reached a higher stage of progress.

the ostracoderms had a cartilaginous skeleton and were without true jaws and paired fins.) Restoration of Cephalaspis. dorsal view. (From Beecher. 180). Figure 248 shows an ostracoderm belonging to a later geologic period. Length about 6 inches. were not common. There is no record of land plants in the Ordovician deposits.) the cyclostomes (see p. twice natural size.362 FUNDAMENTALS OF BIOLOGY derma and. a Devonian ostracoderm. 247. to which they are related. 248. "Evolution of the Vertebrates and Their Kin. B. . restored. FIG. Blakiston's Son & Company. Like A FIG. side view. by permission. ventral view. B An Ordovician trilobite (Triarthrus becki). It has not been determined from whicji invertebrate group the first vertebrates arose. A. but it is thought that pteridophytes were in existence then because of their abundance during subsequent periods." P. (From Patten.

Some of them reached a length of The sharks of the Devonian were also abundant and some of them being 6 feet long. displaying a great variety of types. time (Fig. period when the fishes became the dominant animal group. from the Length 8 Dinichthys. these representing the oldest known land plants. Like the ostracoderms. but the remains of a few primitive pteridophytes have been found. marine invertebrates were abundant. rose into prominence.) The ostracoderms soon be'came extinct. derms (particularly crinoids) made a great notadisplay. (Courtesy of American Museum of Natural History. while sharks. The trilobites. the brachiopods maintaining their numerical The cephalopods superiority. . were more complex and remained powerful. The arthrodirans were the most numerous and the largest ransj fishes of the 20 feet. The Devonian marks the FIG. they had external armor and a cartilaginous internal skeleton. Other primitive types. powerful. presented Other a bizarre appearance. representing a distinctly higher type of fishes. but the arthrodiwhich possibly were descended from them. A the oldest animals. During the Silurian. 249). 249. which culminated in the Ordovician. made their appearance. ble feature of the Silurian was the appearance of scorpions. ostracoderms became abundant.THE LIFE OF THE PAST 363 Silurian and Devonian. but were gradually being displaced by Corals and echinothe fishes. an arthrodirau Devonian shales of Ohio. were reduced in number of species by approxi- mately one-half. by the development spines and tubercles. These early sharks were very primitive and not common. sist known air-breathing Plant remains conmainly of algae. Some of of them. feet.

Restoration of Devonian fishes from the Old Red Sandstone of an ostracoderm. however. 5. myriapods. existence consists of a single three-toed footprint found in Pennsylvania in a late Devonian deposit. have been found in Lower Carboniferous (Mississippian) deposits. 3 and 4. Physical conditions during the late Silurian and early Devonian were favorable for the origin of land animals. but land vertebrates were also present. The land plants of the early and middle Devonian were chiefly primitive pteridophytes like those of the Silurian. while the trilobites declined still further. appropriately called "seed ferns/ Carboniferous and Permian. 2. recognizable with certainty as such.) 1.and primitive gymnosperms. the only evidence of their FIG. sharks. so far as known. The fishes were still a 7 . brates. The oldest amphibian remains. Some of the latter were fern-like plants with seeds. lungfishes. (Courtesy of American Museum of Natural History. and were mostly of small simple types somewhat resembling modern salamanders. In fact. ganoids. 6. Amphibians were not abundant at this time. included scorpions.364 FUNDAMENTALS OF BIOLOGY such as the ganoids and lungfishes. 8. but there were no representatives of the teleosts (bony fishes). arthrodirans. although very rare. 250). but there is now evidence that they were derived from primitive fringe-finned ganoids. 250. while those of the later Devonian were advanced types including both pinMophvie. may have furnished a stimulus that resulted in their appearance. It was once thought that the first land vertebrates evolved from ancient lungfishes. and snails. The air-breathing invertebrates of the Devonian. made their appearance (Fig. and 7. Increasing aridity. causing a drying up of lakes and streams. Scotland. the great modern group that includes over 95 per cent of all living Brachiopods were the most abundant marine invertespecies.

5. " Henry Holt & Company.) Salisbury. for here . some with curious proportions. In the Upper Carboniferous (IVnnsylvanian) the amphibians were the dominant animal group and reached the highest point A composite group of leading plants of the Upper Carboniferous. a primitive gymnosperm. 3. 1. these Carboniferous forms were gigantic. but the sharks reached Most of them were ancient their point of greatest abundance. The earliest known insects Carboniferous age. in their development. Compared to modern insects. Cordaitfs. Lepidodendron and 2. Catamites. a seed fern. gigantic lycopods. Many peculiar types were evolved. "College Geology FIG. such as a small body with a large flat head. by permission. The arthrodirans became extinct early in the period. but this group must have origi- we find an abundance of them and a marked development of the primitive orders. 251.THE LIFE OF THE PAST 365 mighty group during the Lower Carboniferous. nated earlier in the Paleozoic. Sigillaria. but many are so are of Upper cfosely similar to amphibians to recognize them. In the Upper Carboniferous strata primitive reptiles that it is difficult have been found. a horsetail. shell-feeding types that later became almost extinct. (From Chamberlin and 4.

of Primitive reptiles them were still amphibian-like. but most many forms The and of life were unable to survive into the next era. the event of greatest magnitude was the formation of the Appalachian Mountain System. forests coupling chiefly of gigantic lycopods and horseas well as seed ferns and other primitive gymnosperms Some of these plants reached a height of 100 feet. scorpions. No larger insects have ever lived.) Although the amphibians passed their climax in the Upper Carboniferous. 251).366 FUNDAMENTALS OF BIOLOGY Cockroaches 4 inches long were not rare. The Permian is the last period of the Paleozoic era. 252. the older pteridophyte types declining and gymnosperms becoming more highly developed. Length about 10 (Courtesy of American an amphibian from the Permian of Texas. The vegetation underwent marked changes. It was during this period that our most extensive coal deposits were laid down. moist. Some reached a length of 10 feet (Fig. Fio. (Fig. flies swamp tails. and toward its close great geologic changes that exerted a profound influence upon life took place. Spiders. Widespread aridity and glaciation occurred during the Permian. In North America. the . myriapods. Skeleton of Eryops. THE MESOZOIC ERA close of the Paleozoic marks the decline of the amphibians the beginning of reptilian ascendancy. A warm. Museum of Natural History. while some of the dragonwere 15 inches long and had a wingspread of 30 inches. uniform climate prevailed throughout the Upper Carboniferous. Coal represents accumulated plant remains subsequently metamorphosed by Over immense areas there extended great geologic agencies. making conditions so severe that became common. and this favored the development of a most luxuriant vegetation. 252). they were still abundant in the early Permian. feet. and snails were also common on land. The Mesozoic era.

such as dinosaurs. ichthyosaurs. True ferns.THE LIFE OF THE PAST duration of which " is 367 estimated at 135 million years. Jurassic. The Mesozoic includes four periods the Triassic. the reptiles reached the height of their ascendancy. like modern monotremes and probably insecThese little tivorous in their habits. the land. for example. of Some gymnosperms. and continuing through the Cretaceous. Lower ately called the Age : Cretaceous lived over and Upper Cretaceous (Fig. became more abundant. and the air. creatures were very insig- In fact. Beginning with the Jurassic. their subordinate position doubtless being due to the supremacy of the reptiles. Life of the Triassic. The trees in the petrified forests of Arizona. which are of early Triassic age. others like conifers. From a few simple reptilian types that from the Permian into the Triassic. there arose many new saurs. but a few persisted into the early Triassic. dominating all other animal groups. They dominated the sea. Life of the Jurassic. 246). The appearance of primitive mammals was a notable feature ichthyosaurs of the Triassic. as well as the seed ferns and other primitive gymno- sperms of the Paleozoic. which were the domiof these gymnosperms were much like modern cycads. The first dinosaurs were of small or medium size. however. they were rare in the Triassic and but little specialized for marine life. The ichthyosaurs. The perhaps somewhat nificant They were small. are all conifers (Fig. egg-laying forms. characteristic land animals of the Mesozoic. had become extinct before the close of the Permian. reptile-like. little evolutionary progress. thousands of their footprints are preserved in the Triassic sandstones of the ConThe dinosaurs became the most abundant and necticut Valley. pterosaurs. 241). is appropriof Reptiles/' for at that time they reached their greatest display. the most highly specialized of marine reptiles. and relatively unspecialized. mammals remained comparatively rare throughout the Mesozoic and made members of the Triassic fauna. groups. Nearly all the giant lycopods and horsetails. many ways the most remarkable of and plesiosaurs were lung-breathing aquatic reptiles descended from terrestrial ancestors. The pterosaurs all were flying forms and in reptiles. lizard-like. plesioand turtles. attained their climax early in the Jurassic. The plant life of the Triassic was characterized by the appearance of new and higher types nant plant group. They were .

368 FUNDAMENTALS OF BIOLOGY distinctly fish-like in appearance. feet. 253). (Courtesy of American Museum FIG. a short neck. The iohthyosaurs were viviparous. of Natural History. a highly specialized marine reptile from the Jurassic Length 25 to 30 feet. 253. the figure represents a mother with brood of young. Germany. a dorsal and a caudal fin.) armed with numerous sharp teeth ichthyosaurs attained a length of 30 lizards (Fig. which by the end of the Jurassic reached their cul- . and a slender pointed snout of Ichthyosaurus. Some of the Crocodiles and true made their first appearance in the Jurassic. with paddle-like limbs. The most numerous and most powerful land reptiles were the dinosaurs.

sharp.) long. there then being the greatest variety and the largest ones that ever existed. The creature is shown preying upon one of its herbivorous Allosaurus lived in North contemporaries. All the digits were provided with large. (Courtesy of American Museum of Natural History. 254. often 3 feet in length. but the larger ones walked. Length of specimen 34 feet. 254). The jaws.THE LIFE OF THE PAST 369 mination. on . Some of the smaller herbivorous dinosaurs all were bipedal. Skeleton of the bipedal carnivorous dinosaur Allosaurus and restoration of the same. America during late Jurassic and early Cretaceous times. but during the Jurassic both carnivorous and herbivorous types were common. like all the carnivorous forms. with hind legs much larger than the fore legs and alone used in walking (Fig. bore numerous sharp teeth. the former preying upon the These flesh-eaters were powerful creatures 30 to 35 feet latter. Nearly all the Triassie dinosaurs were carnivorous. FIG. curved claws.

255. These huge beast s were sluggish in their movements and probably lived in swamps. . (Courtesy of American Museum of Natural History.) but the body was less bulky and the tail considerably longer. the neck and tail long. quadrupedal. herbivorous dinosaur from the late Jurassic of Wyoming. a gigantic. Length 66 feet. In all these herbivorous giants the head was comparatively The toes small. This creature inhabited swampy meadows and flood plains. and the legs massive. No larger land animals have ever inhabited the earth. Diplodocus reached a length of 80 to 90 feet. One of the largest of these creatures was BrontosauruSj with a length of nearly 70 feet and an estimated weight of 37 tons (Fig.370 FUNDAMENTALS OF BIOLOGY four legs. were short and the claws blunt. Brontosaurus. and estimated weight 37 tons. 255). FIG.

Length about 20 feet. The pterosaurs made progress during the Jurassic and became more numerous. 256. with narrow bodies having great plates projecting upward from the back and long tail. The head. The oldest fossil birds are known from the Jurassic. (Courtesy of American Museum of Natural History. did not evolve from pterosaurs. Stegosaurus must have been extremely stupid and sluggish. but were supported by only one elongated digit. but from some more remote reptilian stock. They . relying upon its armor for its more active flesh-eating contemporaries. stretched from the fore limbs to the hind limbs and body. and some had rudder-like tails. Stegosaurus. a quadrupedal herbivorous form of Jurassic age. 257). reached a length of about 20 feet (Fig. Restoration of the armored dinosaur Stegosaurus.THE LIFE OF THE PAST Another group of 371 quadrupedal Jurassic dinosaurs of herbivo- rous habits were the grotesque armored forms. possibly in the Permian. The animal was about the size birds a true connecting link. 256). but the group must have originated earlier. one of the best known of these dinosaurs. as the structure of their wings clearly shows.) 3 feet. In the lithographic limestone of Bavaria two complete skeletons have been found of a very primitive bird called Archaeopleryx It is clearly a transitional form between reptiles and (Fig. which was unusually small. The largest ones had a wingspread of about protection against FIG. Most of these flying reptiles had bird-like heads with The wings teeth in their jaws. as in modern bats. had beak-like jaws and a tiny spines on the brain that weighed about 2J/ ounces.

clawed digits. separate pelvic bones. and teeth in both jaws. Restoration of Archaeopteryx. a primitive Jurassic bird showing " X Js- (From Romanes.'"' Open Court Publishing Company.) conspicuous reptilian characters were a long vertebrated tail. wings with three free. . its FUNDAMENTALS Its OF BIOLOGY most head and the presence distinctive avian features were the shape of of wings and feathers. by permission. many reptilian features. Darwin and after Darwin . Some of its FIG. 257.372 of a crow.

the extreme specialization and final extinction of the great reptiles. or was the appearance bony fishes. Tyrannosaurus. but shortly their place was taken by other groups. 258). a highly specialized carnivorous dinosaur attacking Both forms lived during the Upper Cretaceous. a horned herbivore. Of the dinosaurs that evolved during the Jurassic. nearly 50 feet in length and with jaws 4 feet long. of the gigantic herbivorous dinosaurs became extinct before or soon after the beginning of the Cretaceous. size. Before the end of the Mesozoic they posi- The life of the Cretaceous presents two important developments. Life of the Cretaceous. viz. a tion which they have held ever since. and ferocity the most destructive life engine which has ever evolved" (Osborn). had become the dominant group of aquatic vertebrates. FIG.) the large carnivorous forms lived well into the Cretaceous but finally died out and were succeeded by huge specialized creatures that surpassed even them in ferocity. This terrible beast. THE LIFE OF THE PAST of the Jurassic 373 of the Another feature teleosts. Triceratops. (Courtesy of American Museum of Natural History. was "in respect to speed. Of the Upper Cretaceous flesh eaters. were bipedal forms with webbed feet and a long flat .. power. the duckbill Most dinosaurs. 258.. and the appearance of angiosperms. One of these. the most notable was Tyrannosaurus (Fig.

One of them (Pteranodon) with a small body. Length about 25 feet. 260). FIG. very bulky. head. Like the dinosaurs. 259.-proad of 20 feet (Fig.) were present. It was the largest and most highly specialized flying creature that has ever lived. these were used for grinding vegetable food.374 tail. 258). had a \\mg. the latter of defense. Like modern birds. These curious dinosaurs were 20 to 25 of the Upper Cretaceous. The horned dinosaurs were among the last to become extinct at the close of the Mesozoic. Mighty battles must have been waged between Tyrannosaurus and Triceratops. most of the Upper Cretaceous also . and feet long. Restoration of the duckbill dinosaur Trachodon. quadrupedal (Fig. As in the tops. a bipedal herbivorous form of Upper Cretaceous age. The head and the limbs short but mason the Horns were present sive. FUNDAMENTALS OF BIOLOGY they fed upon the vegetation of swamps and marshes (Fig. three of them in Tricerafrill occurred over the neck. . (Courtesy of American Museum of Natural History. Another group of herbivores were the horned dinosaurs 259). the jaws were beaked and only back teeth was large the tail relatively short . and a massive bony duckbill dinosaurs. the flying reptiles of the Upper Cretaceous became highly specialized. the former representing the climax of dinosaurian offense.

so far as known. In fact. the head being smaller and the neck longer. oaks. The evolution of the first angiosperms was so remarkably rapid that by Upper Cretaceous times they had spread over the entire earth. mosasaurs were also abundant at this time. such familiar trees as willows. 260. dominating the vegetation and giving it a distinctly modern aspect. cycads. the ichthyosaurs declined rapidly during the Upper Cretaceous. large forms incapable of flight. History. In addition to marine turtles and crocodiles. But with the appearance of the angiosperms later in the period. over half of which was neck. and i-n'Mf. were specialized for diving. Of the aquatic reptiles. an Upper Cretaceous pterosaur with a (Courtesy of American Museum of Natural nearly 20 feet. carnivorous. The birds of the Cretaceous period. lizards. while the plesiosaurs reached their culmination (Fig. but the largest of the Cretaceous forms (Elasmosaurus) exceeded 40 feet in length. and all having teeth. these ancient types declined. marine length of 40 feet (Fig. 261). some reaching a During the early part of the" Lower Cretaceous the flora con- sisted of ferns. plesiosaurs were not much over 20 feet long.) wingspread of Restoration of Pteranodon. The latter were less fish-like than the The Jurassic former.-rji-ii!i|. . 262). They were gigantic.that had flourished during the Triassic and Jurassic.THE LIFE OF THE PAST 375 pterosaurs were short tailed and toothless. some being FIG.

) . by permission. (Courtesy of chasing the giant the Upper Cretaceous of Kansas. FIG 261. fish Portheus. American Museum of Natural History. a mosasaur from These sea lizards attained a lengih ot 40 feet. Length about 10 feet. (From WiUiston.) FIG 262. " the Past and Prevent. TVoaoMrwa. Water Reptiles of University of Chicago Press. Restoration of Trinacromerum.376 FUNDAMENTALS OF BIOLOGY an Upper Cretaceous plesiowuir.

It is characterized as the "Age of Mammals " and of life. and crocodiles members of the modern fauna. As already noted. leaving to represent them today only the lizards. after holding sway for approximately 135 million years. With the earth. and the Quaternary. 377 tulip trees. and palms were common. all insignificant THE CENOZOIC ERA Approximately 60 million years have elapsed since the beginning of the Cenozoic. snakes. the duration of which is therefore less than half that of the Mesozoic. sassafras. primitive egg-laying arose early in the Mesozoic but remained rare and . which terminated approximately 1 million years ago. The close of the Mesozoic was characterized by stupendous geologic changes that took their toll of life. But. viously taken place at the end of the Paleozoic. just as had preFor example. great extinctions of radical changes in environment going on all over Just what factors life occurred.THE LIFE OF THE PAST elme. The represents the most recent chapter in the history Cenozoic includes two periods of very unequal length: the Tertiary. were responsible for bringing about the ultimate extinction of the great Mesozoic reptiles may perhaps never be known. the period in which we are now living. maples. turtles. they did disappear from the earth. magnolias. figs. The with the ensuing discussion of following table will be useful for reference in connection mammalian evolution: mammals Life of the Tertiary. the formation of the Rocky Mountain System occurred at this time.

These archaic Their feet were pcntadactyl and mainly plantigrade. period. . These gradually displaced most of the archaic forms. have been a steadily declining group ever Birds were probably numerous throughout the Tertiary. At the beginning of the Eocene there appeared in both Europe and North America. bats. and some were present that have no modern representatives. the During their climax during the late Miocene or early Pliocene. as invaders from another part of the world. while even Greenland had a rich temperate flora. mammals. vailed in most parts of the world during the early part of the In fact. Nearly all of them had the primitive number of teeth (44). The later Tertiary was distinctly cooler. however. Nearly all the modem orders were differentiated at this time. as they are called.378 FUNDAMENTALS OF BIOLOGY At the close of the little progress throughout the era. while the Quaternary marked the advent of the Great Ice Age. forms. which developed in two sets but showed little differentiation. laurels. This adaptive mammals continued throughout the Oligocene. in the Eocene such tropical plants as figs. palms. included some groups that soon became extinct but also others (such as inseetivores. particularly A warm moist climate preof the younger herbaceous groups. Adaptation to different modes of life radiation of resulted in the development of diverse types. reaching since. magnolias. and primates) which gave rise to modified descendants that still live. with small brains but far more intelligence than any of the reptiles. mammals of more advanced types. is known regarding their fossil The vegetation of the Tertiary was characterized by the great abundance and rapid evolution of the angiosperms. Miocene the mammals made a remarkable display. except for man. when many new forms arose more nearly like those of the present. and breadfruit trees grew in parts of Europe and North America now temperate. Mesozoic. true placental forms being found in the Paleocene These early Tertiary mammals were small generalized beds. as they are today. but little history. Mammalian ascendancy began early in the Tertiary. the extinction of the great reptiles gave these ancestral mammals an opportunity of which they took made immediate advantage. and.

wolves. beavers. camels. The oncoming of each successive glacial advance resulted in wholesale migrations and extinctions of life. Human evolution forms the subject of Chap. mammoths. gressed along two general lines: for rapid running over hard ground. camels. In North America five distinct glacial invasions took place. while temperate forms retreated to ning of the more genial regions. and elephants. tapirs. There is has Quaternary period conclusive proof that man lived in Europe throughout the greater is no undisputed evidence of North America during this time. an abundant and varied fauna occupied temperate North America. since then about 25. Adaptive Features of Modern Horses. XXIII. During the warm interglacial stages. but there his existence in EVOLUTION OF THE HORSE AND ELEPHANT Because the Cenozoic deposits have been formed during relamuch has been learned concerning the life of The evolution of many mammalian groups has been this era. bison. A notable collection of Pleistocene mammals has been obtained from the asphalt pits of southern California (Fig. tively recent times. thoroughly traced. Thus the legs . The most outstanding event part of the Pleistocene. separated by long intervening warm stages. The evolution of horses and elephants is so well known that no account of paleontology would be complete without a brief consideration of their fossil history.000 years have elapsed. Arctic animals and plants came southward. and saber-toothed cats. deer. horses. Modern horses and those of the Pleistocene belong to the genus Equus. 240). in the history of life during the been the evolution of man. and for grazing. The begin- Recent period dates from the withdrawal of the last ice sheet. examples being the carnivores.THE LIFE OF THE PAST Life of the Quaternary. mastodons. rhinoceroses. bears. The structure of the horse has been modified in accordance with these functions. peccaries. horses. tapirs. antelopes. 379 feature of this last The most notable period in geologic history was the extensive glaciation that occurred during the Pleistocene as a result of great climatic changes. pigs. including giant ground sloths. ruminants. a highly Specialization has prospecialized branch of ungulate stock.

reduced in the male. It had a 44 of teeth. both the skull and The elongated skull. neck. the former being very small and the latter large. lived in The most the masticating surfaces having a primitive pattern. In order to permit the head to reach the ground without the necessity of the animal's bending its knees. descendant of a five-toed ancestor. as well as those of the shank. The evolution of the horse took place chiefly in North America. . There was no trace of the first digit on the fore feet. flat. The wrist and heel do not touch the ground. It has been traced through a number of successive stages. The canines are greatly* teeth. 213C). with the neck have become lengthened. but on the hind feet vestiges of both the first and fifth digits were Thus there can be no doubt that Eohippus was the present. and limbs height. the hind limbs three. and modified for grinding. some of which will now be considered. Eohippus was a slender animal about a foot in Its head. were relatively short and the two bones comprising the forearm. functional toe and that corresponds to the middle digit of a pentadactyl limb. " of two slender splint bones. primitive genus of horses is Eohippus. a form that North America and Europe during early Eocene times (Fig. also provides room for the development of The incisors. being broad. the foot consisting of a single. while in the hind limb the fibula has similarly united with the tibia. 263). and usually are entirely absent in the female. They are very high crowned and are provided with a complex masticating surThe grinding teeth continue for a long time to grow in face. Stages in Equine Evolution. height as they are worn down. the fore limb. this being provided with a large hoof. 263). are very large and strong." representing the remnants of The horse walks upon the second and fourth digits (Fig. The fore limbs of Eohippus bore four functional toes.380 FUNDAMENTALS OF BIOLOGY IE are very long and slender and can bend only in one plane. Modern the tip of the too. The premolars and molars are similar in form and function. or about the size of a fox. enlarged. the ulna has become greatly reduced in size and has fused with the enlarged radius. horses are about 5 feet or more in height. were not yet fused. The set teeth were low complete grinding crowned. which function as cropping the teeth (Fig. its large deep jaWvS.

History.THE LIFE OF THE PAST 381 FIG. 263. (Courtesy of American Museum of Natural .) Skull of pus) photographed to modern horse (Equus) and model of Eocene horse (Eohipsame scale.

but the third toe on both the fore and hind feet was larger and the vestigial digits had dis- The second horse appeared from the latter. to a grazing habit. or about the size of a Its head. being 3 to 4 feet in height. All the feet were three toed. Pliohippus was. but more complex than those of the Eocene horses. and Africa but for some unknown reason became extinct on the American continents and in Europe during . while the two lateral ones did not touch the ground. like Merychippus.382 FUNDAMENTALS OF BIOLOGY in our series is Orohippus. It was considerably larger than its forerunners. while forms capable of becoming adapted the Oligocene. the middle one being the largest. Each foot was provided with three functional toes. neck. whose remains have been found in the middle Eocerie^leposits of North America. Most of the browsing types of mammals became extinct in North America at this time. progenitors. Mesohippus was a North American horse that lived during It was 18 to 24 inches tall. Pliohippus gave rise to Equus. It was only slightly larger than Eohippus. and limbs were longer than those of its sheep. involving an extensive development of prairies and areas suitable for grazing. fibula had become greatly reduced and was partly fused with the tibia. a late Pliocene form. The fore limbs bore a vestige The teeth of Mesohippus were low crowned of the fifth digit. Asia. Merychippus lived in North America during the Miocene. In this horse the radius and ulna were completely fused. not much larger than Merychippus. Europe. but the middle digit was relatively larger than in Mesohippus. The two lateral toes were reduced to splint bones. The teeth of Orohippus were but slightly more advanced than those of its forerunner. which spread from North America to South America. but they were incapable of continued growth. but the skull was still relatively short and uuspccialThe ulna was very slender but still distinct. while the teeth were more highly crowned and had complex grinding surfaces. The grinding teeth were relatively high crowned and their surfaces somewhat complex. Merychippus is thought to have been the first horse to have turned from a browsing to a grazing habit. and a disappearance of the older forested regions. but advanced in other ways. The Miocene was an epoch of continental elevation. and the number of functional digits was the same. flourished. Its skull was longer and its lower jaw heavier. while the ized.


Adaptive Features of includes two modern from the Modern Elephants. Some of the mammoths of the Pleistocene were as large or slightly larger than this (Fig. The genus Elephas species of elephants and the extinct mam- FIG. in addition to these six studied. 265. Knight. but living elephants occur only in Asia and Africa. while the largest African elephants are said to be nearly 13 feet tall. and North America. The Indian species attains a maximum height of 10 feet. The limbs of elephants are large and pillar-like. a number of transitional forms have been found in inter- mediate strata. Asia. Restoration of imperial mammoth (Elephas imperator) by Charles R. Figure 264 graphically presents the evolution of the horse jbhrough the six stages described above and should be carefully It must be understood that. giving an unbroken evolutionary series early Eocene to the present time.384 FUNDAMENTALS OF BIOLOGY the Pleistocene.) moths. During the Pleistocene the genus was widely distributed over Europe. surviving only in parts of Africa and Central Asia. elephants are the largest mammals. stages. 265). living With the exception of the whales. an obvious adaptation for . (Courtesy of American Museum of Natural History.

forward in the jaw. had a higher skull. it the only teeth that the elephant has are molars. The chewing surface is very complex. The neck was long enough to have enabled the head of the animal to reach the ground.THE LIFE OF THE PAST 385 supporting the enormous weight of the body. and grinding teeth with two or three transverse ridges. and of these never has more than four complete or eight partially worn ones As the grinding teeth are worn down. The tusks are highly modified incisor teeth that reach an extreme length of 8 feet in the Indian elephant and 10 feet in the African species. large air spaces being also present. The skull of the elephant is very short and high. 266). The molar teeth were larger than in the earlier form. The feet are five In most long-legged mammals the toed. The brain is large nearly twice as large as the human brain but it does not fill the cranium. The dentition of the elephant is highly specialized. The trunk is a prolongation of the nose and upper lip. There has been found in the upper Eocene and lower Oligocene deposits of Egypt a mammal genitor of called Moeritherium. or about the size of a pig. It was about 8^2 feet in height. each toe being hoofed. being entirely out of proportion to its length. and longer tusks. The upper lip was slightly elongated and probably reached to the tip of the tusks. but the upper lip may have been prehensile. Stages in Evolution of the Elephant. but the lower tusks were much shorter than the upper ones. in the Indian elephant and the extinct Siberian mammoth. It was larger than Moeritherium. in some of the mammoths the tusks were 16 feet long. It did not look like an elephant except in having a high skull. but in elephants the massive head makes this arrangement impossible. having in the African elephant up to 10 or 11 transverse ridges. two short tusks in each It jaw. a relatively shorter neck. up to 27. and are replaced by new ones appearing behind. had no trunk. Accordingly the trunk takes the place of a long neck. generally recognized as the promodern elephants (Fig. Aside from the tusks. form called Paleomastodon has been found in the lower A Oligocene formations of Egypt and India. it is used chiefly in conveying food to the mouth. they move at a time. . and the three transverse ridges were more conspicuous. neck is also elongated to permit the head to reach the ground.

Trilophodon lived during the Miocene in Europe. arid It was a large animal almost as tall as the FIG." The Macmillan Company. E. F. (From Lull. Pliocene. Stegodon. by permission.386 FUNDAMENTALS OF BIOLOGY North America. ''Organic Evolution. . D' Trilophodon. F'. Eocene. Mastodon. A'. Moeritherium. Pleistocene. Africa. Paleomastodon. Pliocene. C". Evolution of the head and molar teeth of elephants. D. 266. The latter were large and reduced to the same number as in modern modern Indian elephant. C.) t It made an advance over Paleomastodon in having longer tusks and more complex molars. B. Miocene. Elcphas. E'. Oligocene. A.

FHE LIFE OF THE PAST 387 still only three transverse ridges. A The there never being more than eight molars present at any time. and about the size of the modern Indian elephant. but there were modern species. these were about 9 feet in length. transverse ridges did not exceed five or six. of which several species are known.) Restoration. 267. being FIG. CONCLUSIONS The facts of paleontology justify the drawing of certain general conclusions with reference to the course of organic evolution. The lower shortened and the teeth in was reduced greatly jaw number. Trilophodon was peculiar in that its lower jaw was greatly elongated and provided with a pair of tusks. These may be briefly stated as follows: (1) Throughout geologic history there has been an ascending succession of plant and . three being the common number. Kidgway. the far as did the tusks. trunk was short but probably reached as Mastodon. lived during the Pliocene and Pleistocene in North America. The mastodons were larger than the earlier forms. 267). (Courtesy of Los Angeles American mastodon (Mastodon amcricani's) by Museum of History. and Asia. of John L. As compared to elephants. pair of tusks was present only in the upper jaw. but they were stockier in build (Fig. Art. Europe. Science.

. but can change only in the direction (5) of greater specialization. With a radical change in environ- ment. specialized organisms are often unable to survive the now conditions and so become extinct. This is shown by the fossil record of many groups. (2) The of evolution has been toward greater structural trend general specialization in adaptation to particular conditions of living. Thus each succeeding geologic period marks a progressive advance in life. (3) Higher forms of life have sprung from generalized members of lower (4) Forms highly modified groups. for living again become under a particular set of external conditions cannot generalized.388 FUNDAMENTALS OF BIOLOGY in animal groups corresponding to the sequence of rock formations which their remains are embedded. not from specialized ones.

the fact of evolution will always remain. and it has long been assumed that such induced changes are transmitted to subseinfluences 389. So it that. Biologists are unanimous rests Some in their conviction that evolution is a natural process. They are not in agreement. Theories of evolution are usually associated with modern must be borne in mind times. but it should be realized that crude conceptions regarding the derivation of higher organisms from lower ones have existed from the time It has only been of the early Greek philosophers. however. since the beginning of the nineteenth century. however. and consequently a diversity of opinion exists as to what are the most important factors involved. The older evolutionary conceptions were wholly speculative. A number of satisfactory. and it has been seen that all the facts presented in the last two chapters can be explained only on the basis of descent with modification. but none is entirely It is one thing to show that evolution has taken place but another thing to explain how it has taken place. This is because the method by which evolution has come about is imperfectly understood. causative theories have been advanced. that evolution has been the subject of scientific study. INHERITANCE OF ACQUIRED CHARACTERS That organisms can be modified through the action of external is a matter of general observation. but our modern theories of causation are based on careful observation and experimentation. as to how the process operates as to the causes underlying organic changes. .CHAPTER XXII THE CAUSES OF EVOLUTION of the evidence upon which the principle of evolution has been examined. We shall consider only the three most important theories of evolution and take them up in the order in which they have been advanced. although proposed explanations may be discarded.

Jean Baptiste de the cause of evolution was quent generations. Jean Baptiste de Lamarck. 268. Lamarck assumed that the effects produced by a change in external conditions become cumulative through succeeding generations. and higher animals on the other. Lamarck's theory is based on the idea that acquired characters are inherited FIQ. somewhat differently in the case of plants Direct Action of Environment. 314-316). that individual adaptations become racial is that inheritance modified by environment. but function. something imposed upon it by the surroundings. fication arises Lamarck thought that modiand lower animals on the one hand. although in both cases the causal agent is a change in environment.390 FUNDAMENTALS OF BIOLOGY The greatest exponent of this idea as a French naturalist. An a modification that arises as a direct response acquired character to an external stimulus. is whose theory was announced in 1801. in the latter. 268). he thought that racial characters have developed by the inheritance indirectly. such as a change in environment or It is not part of the iirpmi-m'-* inheritance. Examples of structural changes induced by the environment have been given (see pp. Lamarck (Fig. In the former the environment acts directly. . 1744-1829.

for example and yet in such cases members of the . often grobent and tend to tesquely gnarled. Chamberlain. the changes induced in each generation being transmitted to subsequent generations. it must be realized that the power of adaptive response is limited. for example.THE CAUSES OF EVOLUTION 391 of direct responses. the crown becomes very broad and flat. FIG.. the peculiar features of desert plants. Exposed to the sea winds.According to this theory. 269. a tree native lo the central California coast in the vicinity of Monterey Bay. -Monteioy cypiess (Cuprcbvus maciocarpa). Even in the most plastic little organisms the basic features are so firmly fixed by heredity that or no alteration can take place. and the branches. have resulted from the direct action of the desert conditions upon them. (Courtesy of Professor Charles J. Many widely distributed organisms live under a great variety of conditions most com- mon weeds. grow horizontally away from the wind. Only minor adjustments are possible.) Although cases of individual adaptations are often striking.

the horns and hoofs of mammals. atrophy. the tentacles of snails. the webbed feet of ducks and geese. in the organization of individuals everything which has been acquired. Lamarck believed that . basing his belief on the fact that organs are strengthened through use and weakened through disuse. etc. The former then tend to develop.392 FUNDAMENTALS species often OF BIOLOGY same may exhibit relatively slight differences among the other hand. or changed during the course of their life. He thought that the environment does not act directly but indirectly. members of the same species living under a uniform set of conditions may sometimes show considerable variation. as a rule the changes induced are not permanent. . In fact they are said to become "transformed" into alpine species. These call for new habits and modes of life in accordance with which certain organs are used to a greater extent. 269). Among the higher animals Lamarck thought that structural modification arises chiefly through changes in function. the absence of teeth in certain vertebrates. On Another important point should be kept in mind. the progeny often become highly modified. but when their seeds are planted under normal conditions no former environment remains (Fig. As a result of disuse he explained the limbless condition of snakes. It is in this feature of Lamarck's theory that the greatest weak- ness lies. others to a lesser extent. But when taken back to the lowland. When organisms are placed in a new environment. Lamarck explained the development of a great many structures as a result of the inherited effects of use. the latter to effect of the Use and Disuse. even after many generations of exposure to alpine conditions. the degenerate eyes of the mole. Trees indigenous to seacoasts are often highly modified by wind action. they return to their original state. for example. themselves. . and the long neck of the giraffe. impressed upon. for when put back in the old environment the " acquired characters" become lost. a change in external conditions causing an animal to experience new needs. . Thus when seeds or cuttings of lowland plants are grown in alpine regions. the long legs of wading birds. is preserved by generation [heredity] and transmitted to the new individuals which have descended from those which have undergone those changes.

270). external modify the soma must also affect the germ no known mechanism." This means a simultaneous modification of the germ cells by the same influence that affects the soma but without any transfer from the latter to the former. germ cells are set apart early in development from undifferentiated embryonic cells and are not derived from specialized somatic tissues (see pp. Thus. although Lamarck's theory seems plausible and is supported by some modern biologists. So. This means that any induced change undergone by the soma has no racial effect because somatic cells do not become part of the There is fications may next generation. the inheritance of acquired characters is impossible. is that the theory is unsupported by experimental work. disease. mutilation. however. In conclusion. August Weismann. it is rejected by many because it rests upon an extremely unlikely assumption that is not supported by experimental work. A more fundamental objection. NATURAL SELECTION The Darwin evolution greatest name associated with the principle of organic is that of the famous English naturalist. Darwin did two things: (1) He accumulated . Charles (Fig. He held that. 170). to cause a change in the next generation. Weismann influences that later cells. whereby modibe transferred from somatic tissues to gametes. whose views have largely influenced modern opinion. however. or directly by the environment. 229-230 and Fig. Changes in organisms can be produced by use and disuse. Although it is true that the theory of the direct transmission of somatic modifications does explain the development of a great many structures. there " seems to be a permanent effect through parallel induction. many of some other basis. but even after the causal agent has been operative for a great many generations. the induced effect disappears as soon as the cause is removed.THE CAUSES OF EVOLUTION The idea of the inheritance of acquired characters was strongly opposed in 1883 by the German zoologist. it may be said that. as these are the sole means of transmission between parent and Probably in all metazoans except the very lowest. according to Weismann. however. In a few cases. claimed. characters must arise in the germ cells. offspring. both progressive these cases can be explained on and retrogressive. to be inherited.

were published in 1859 in "The Origin of Species/' a book that has had a more profound influence on human thought than any other scientific work ever written. 1809-1882. most plants many more than 10 seeds a year some tens or even . His conclusions. it applies to all living things. (2) he proposed a theory. Of course. to explain how evolution has taken place. (From University Magazine. that of natural selection. Overproduction. Charles Darwin. In 1798.394 FUNDAMENTALS OF BIOLOGY an overwhelming mass of evidence in support of organic evolution and thus proved its reality. FIG. at the end of 10 years there will be one that billion produce descendants of the original plant.) Photo- is itself Darwin's theory is based on several well-established facts and merely a logical inference drawn from them. graph by Leonard Darwin. If an annual plant produces only 10 seeds. and if this rate continues in each generation and all the progeny live. It was this book that resulted in the establishment of the principle of evolution as a fundamental scientific generalization. 270. Mai thus published an essay in which he showed that human populations tend to increase in geometrical ratio a rate greatly in excess of their means of Darwin became impressed with this fact and saw sustenance. based on over 25 years of investigation and study.

a single sturgeon may lay 2 million eggs a year. while most of the seedlings that do develop never reach maturity because of competition with other plants for light. known competition is so severe that only enough individuals survive to Each species has established an equilibrium replace the parents. Such competition may not necessarily involve a struggle to kill an active combat for it may be entirely passive. When man disturbs the balance of nature by killing off certain species. if all the individuals produced were to live. its own set of environmental conditions. even with an enormously high rate of increase. a cod 6 million. it is perfectly true that the normal rate of reproduction in most species is of such magnitude that. Competition results in an equilibrium of species. The fact of variation to differ from other members of their species the tendency of individuals is of the utmost . In other words. an oyster 60 million! Although these cases are extreme. The seeds of most plants. and other vital necessities. only a few of the countless individuals brought into existence can possibly live to maturity. or merely a struggle against adverse physical conditions of the environment. Most animals are destroyed while in a very young This is stage of development. for food and other necessities of life. For example. do not even get a chance to sprout.THE CAUSES OF EVOLUTION 395 hundreds of thousands. called by Darwin "the struggle for existence/' which acts as a check to the high rate of increase. It is a wellfact that. falling into situations unfavorable for growth. or by under introducing foreign species to compete with the native ones. in only a few generations the earth enough to hold them. organisms compete for an opportunity to live. would not be large Competition. moisture. approximately the same number of individuals of the same kind may be found in any given region from year to year. many while still within the egg. It may involve a struggle for room. Consequently there follows a severe competition. Variation. Similarly the number of young produced by some animals is enormous. disastrous consequences often follow. and this remains constant so long as disturbing factors do not arise. This means that. why oviparous species must produce so many more offspring than viviparous ones. where natural conditions remain undisturbed. Because of the limitations of available space and food.

consequently are the ones to survive and to transmit their favorable variations to their offspring. process This Darwin called "artificial selection. Only those individuals varying in the direction of greater fitness would leave offspring. this selective process is assumed to result in a gradual but steady modification of the characters of the species in the direction of better fitness to the environment. variable." Under a constant set of environmental conditions. some may be better adapted to their environment than These individuals have an advantage over the rest and others. Thus the best-adapted individuals are "selected" to survive and to leave progeny.396 FUNDAMENTALS OF BIOLOGY importance. new standards of selectioi\are instituted and evolution is more rapid. . because the fittest. but. namely. individuals are exactly alike. Continuing generation after generation. the selection of certain individuals as the basis for the next generastruggle for life" tion and the elimination of the others. but merely took the fact of variation for granted. more individuals come into existence than can possibly a severe competition in which only a few Because all members of any given species are highly survive. and this leads to of the others perish. the selected individuals being those showing variations in some desired direction. although it often requires examination and a thorough acquaintance with the organism to detect differences. evolution by natural selection is gradual. when conditions change. organisms adapted to the old conditions would not be able to survive under the new ones. Darwin did not distinguish clearly between heritable and non-heritable variations. Survival of the Fittest. This idea of "the preservation of the favored races in the was called by Herbert Spencer the survival of Darwin called the process natural selection. Thus in a region undergoing a 'slight but persistent change in climate over a long period of time. and thus in time many new forms would come into existence and many old ones disappear. while all Infinitely live. and assumed that critical No two practically is all organic differences are heritable. it was suggested to him after making an intensive study of the method used by man in the development of his numerous races of cultivated plants and domesticated animals. Darwin's theory of species formation a logical inference based on the facts that have been presented. for without variation there could be no evolution.

evolution utility. that is. The fact that the theory has certain weaknesses was realized by Darwin In fact. not just adaptive ones. shall consider here some of the most serious limitations of natural We selection as a complete explanation of the origin of species. for example. On the basis of natural selection. of variations. that. and perhaps it is the most satisfactory explanation yet proposed of the origin of adaptive characters. non-adaptive characters that apparently have no direct survival value. lived during the Pleistocene. has gone so far that the individual is really handicapped in the The extinct Irish elk. It ters are useful. however. 397 Granting that a selective process does occur in nature and that the fittest survive. It has been pointed out. which struggle for existence. it is difficult to understand cases of overspecialization where struc- tures have been developed far beyond their point of greatest In many organisms. another on the basis of slight. should be realized. Non-adaptive Characters. exclusive factor in evolution but merely the chief factor.TSE CAUSES OF EVOLUTION Limitations of Natural Selection. but it cannot explain the arrival of the fittest. considerable question has been raised as to how far natural selection is adequate to explain the formation of new species. the theory may explain the survival of the fittest. had horns so large that they must have seriously interfered with its movements. both living and extinct. . he considered natural selection to be not the himself. that not all characnor do they appear ever to have been useful in Most species are distinguished from one the organism's ancestry. it cannot produce new ones. It has long been realized. that natural selection is not concerned with the origin it merely takes the existence of heritable variations . Many paleontologists believe that overspecialization has been a most important factor in causing the extinction of species. Over specialization. however. Natural selection has been used to account for many striking cases of adaptation between organisms and their environment. not seem to give the organism any advantage in the struggle Any complete theory of evolution must account for all characters. trivial. The enormous curved tusks of the mammoths were similarly too highly specialized to have served their original function. although natural selection may be effective in modifying characters already present. Origin of Variations. they do for existence. As commonly stated.

known as Oenothera lamarckiana. selection is amount for the of change. Structures such as wings. tion cannot proceed For example. Not only is natural selection limited in its Incipient Stages. Cumulative Effect of Selection. although a tall tall beyond the natural range of variability. practically all variations are heritable. average but cannot transcend the limits It merely tends to isolate pure of variability that already exist. in an useful. parents tend to produce tall offspring. to be effective. may raise the general MUTATION Vries. FUNDAMENTALS OF BIOLOGY and then explains how they are preserved and modified. horns. strains from a mixed population. operation to characters already present. and race will result from the continued mating of tall no more extreme than Selection individuals. although most of the plants in the field were . selection based on them does not bring about a permanent change in the race and hence can have no evolutionary value. the tallest offspring will be those which occur among the general population. Although Darwin assumed that Heritability of Variations. it has been found that With respect to their origin. but it cannot act upon structures until they have developed sufficiently to have become Otherwise it would give the organism no advantage in the struggle for existence. tions are now somatic variations are not heritable.398 for granted. undeveloped condition they are useless. the latter by the environment. Even when confined to germinal variations. and many others have no survival value until fully formed. Thus selection. two kinds of variarecognized: germinal and somatic. Among able to bring about only a very limited a mixed population. but gradually selection becomes In other words. selecineffective in producing further change. which had been introduced into Holland as a garden plant and was growing wild in the vicinity of Amsterdam. Because many are not. The former are determined by heredity. De Vries observed that. selection in any given direction tends to create pure strains relatively uniform characters involved. must be confined to germinal variations. a The mutation theory was announced in 1901 by Hugo de Dutch botanist. It was based largely on a study of a cultivated species of evening primrose.

for the most part. by permission. and rosettos. of red veins. " Mutationstheorie. short styles. a few were -lrikin. 271.) some always a relatively few new individuals each characterized by striking peculiarity. which de Vries found altogether about 12 different types. came true to type. right). ung. 271). to constitute new species (Fig. and Gruppenweiae ArtbildFIG. These new individuals. mutants. left) and one mutants (0. different enough in fact." Gebruder Borntraeger. and thus their .THE CAUSES OF EVOLUTION 399 typical in every way." 1st cd. He then took seeds from typical lamarckiana plants.rly different. he Further breeding revealed the fact that these called mutants. tall habit. sowed them in his botanical garden. Von Veil & Company. etc. Berlin. and found that among the progeny there were of its " Lamarck's evening primrose (Oenothcra lamarckiana. dwarf habit. gigas. flower stalks. such as smooth leaves. (From de Vrics..

most mutants are not so fit as the species from which they have arisen. If better adapted to the environment than the parent species. but through the struggle for existence natural selection determines whether or not these new forms will survive. that most new varieties have originated in this way. the changes appearing among the individuals of X . Although it seems improbable that environmental factors are concerned with the appearance of mutations. small. forming a graded series from one extreme to the other. and are constant. of numerous. which suggested that new species appear suddenly. but if not so well adapted. but one not so prominent as that ascribed to it by Darwin. intergrading De Vries. in both plants and animals. are distinct from the beginning. that gene mutations may be induced by treatment of germ cells with rays. sidered of prime importance are called discontinuous variations or mutations because they are not connected with each other by intergrades. de Vries proposed the mutation theory. They think that new species come into existence by the spontaneous appearance of mutants. variations under the influence of natural selection. through many generations. considered that new species are formed by the gradual accumula- tion. and under natural conditions would not survive. It should be understood that the factors responsible for the appearance of new characters by mutation are largely unknown. give natural Relation to Natural Selection. The outstanding between the theories of natural selection and mutation Darwin lies in the kind of variations that each emphasizes. In fact. de Vries thought . lllifjjf emphasized are called continuous variations or fluctuations because vary about a standard. they persist. as where a group of men are arranged in a The variations that de Vries conline according to height. it has been discovered. yet the conditions under which they arise are not understood. It seems certain that mutations arise from internal causes operating on the germ cells. thought that new species arise suddenly and are The variations that Darwin distinct from the beginning.400 differences FUNDAMENTALS OF BIOLOGY were constant. Mutants have frequently arisen in plants and animals under domestication (see p. difference Continuous and Discontinuous Variations. they are eliminated. on the contrary. The mutationists selection a role in species formation. 282) in fact. As a result of these studies.

nevertheless the possibility of the hybrid nature of this species exists. former category belong variation and heredity. Thus. even if this ratio does not offer ready explanation according to Mendclian principles. (2) Although the evening primrose studied by de Vries is unquestionably giving rise to new types.THE CAUSES OF EVOLUTION 401 the next generation. In spite of the fact that a great many species have been studied.5 per cent of the progeny of lamarckiana plants. involved. such cases are comparatively rare. fectly are factors and that the problem many cooperating involved. Of the various evolutionary factors. certain. Objections. and. . a complete answer will not be forthcoming until a great deal more knowledge of plants and animals has accumulated. it can have but slight significance as a means of species formation. Such treatment results in mutations greatly in excess of the number that would otherwise appear. however. it must be admitted that most of them are not in a "mutating" condition. Although much progress is being made toward a solution of this great central problem of biology. some are primary or To the causative. if mutation is not a general phenomenon. least observed that mutants appear in about 1. and although it has been found that a number of other plants and animals are behaving similarly. Thus the method is still imperhave which about come changes evolutionary by It seems that several or understood. (1) An objection that at once occurred to de Vries and has been more or less urged ever since is the possibility that Oenothera lamarckiana is a hybrid or at an impure species that the "mutants" are merely segreDe Vries gates arising from the "splitting" of a hybrid. Two important objections have been raised against the mutation theory. while others are secondary or directive. CONCLUSIONS The various theories that have been presented in an attempt how the process of evolution operates represent a difference of opinion as to what are the most important factors to explain have seen that no one theory is entirely adeits strong and weak points. to the latter the environment and natural selection. We quate each has is of explaining the causes underlying racial changes in structure of much greater complexity than the early students of evolu- tion thought it to be.

for each kind may be either germinal or somatic. intensive study of the mechanism of hereditary transmission and has focused attention upon the chromosomes of the germ cells as Whether variations the probable place where heritable variations originate. which are comparatively rare. are continuous or discontinuous is of little consequence.402 FUNDAMENTALS OF BIOLOGY Since it is Causative Factors. . Mutations may be either large or small. Small mutations. sort. Mutations arc heritable variations that are not due to segregation or recombination of characters. It is they which are gradually 1 For example. however. Somatic variations arise from the first cause. 175). are Mutations of this thought by some to give This is the method of evolution rise to new species directly. They apparently arise spontaneously in the germ cells from unknown causes. often involving several characters at once. most of the heritable variations exhibited by the fruit fly (Drosophila) are of this type (Fig. and it is this type of variations that forms the raw material for natural selection to work upon. impossible to determine to which one of these three causes any given variation may be due. that they arise independently The pioneer work of Mendel initiated an of the environment. The former. a knowledge of the causes of variation and their manner of hereditary transmission is fundamental to any complete explanation of evolution. our knowledge of the causes operative in the production of germinal variations is very meager. For this reason much modern research is being directed along these two lines. which are very 1 common. arise mostly as sudden changes involving individual genes. emphasized by de Vries. functioning. they can be distinguished only by breeding. It seems rather certain. germinal variations from On the basis of outward appearance it is usually the other two. Although we have learned to differentiate between somatic and germinal variations and to recognize that it is probably only the latter that are of evolutionary value. are frequently due to the addition or loss of chromosomes in the germ cells arising from irregularities in their distribution at the time of reduction. (3) mutation. The important thing to any to recognize that variations may be due one of three causes: (1) differences in environment or is recombination of hereditary characters or (2) reappearance (segregation) of latent ones. obvious that there could be no evolution without variation.

Although this means that the environment it of Darwin deal with tho the latter is generally but development adaptive characters. So far as the application of the selection principle to the racial improvement of man is concerned. considered to be by far the more adequate. natural selection determines which individuals. and one of the The history great problems of biology is to explain adaptation. It is generally considered to be the most important directive factor yet discovered. Selection is also not a causative factor because it cannot originate variations.. individuals that survive in the struggle for existence are. as revealed by the fossil record. but since it probably acts indirectly it cannot be a causative factor. plainly teaches that there has been a marked correlation between changes in physical surroundings and the appearance of new types of organisms. is an important it does not that is a direct one. determining the course of evolution. that . the latter resulting from spontaneous variability The environment is an important arising from primary causes. or social. According to the Both the theories of of Lamarck and theory of natural selection. other words. provided that they have survival value. shall be preserved. . among a population exhibiting infinite diversity. Directive Factors. of life. the past evolution of the human race has been guided by the elimination of the unfit. directive factor. in general. as Conklin says. whether physical.. it is certainly true. Thus the environment determines survival but not fitness. it merely preserves and accumulates such In as already exist. those best adapted to their environment. intellectual. as evolutionary factor. and the future progress of the race must depend upon this same process. imply Lamarck and other early evolutionists assumed. The general trend of evolution is toward greater fitness to the conditions of existence.THE CAUSES OF EVOLUTION 403 accumulated and built into new species according to the Darwinian conception.

and that it is subject to the same natural laws that apply to other organisms. the baboons. physically they have remained primitive. grows. but in all the higher primates there are 32 (Figs. arboreal in their habits. with the excepinfancy. Primates are largely herbivorous and. the first digit being others. are confined to warm regions. and is nourished in the same way. Primages are plantigrade mammals with pentadactyl limbs. Therefore whatever causes have played a part in the evolution of other animals have also been operative in the derivation of the human species from a more primitive ancestor. Like the limbs. The hands.. and some of the Old World macaques. more or less opposable to the are prehensile. clavicle (collarbone) is in the this is also a primitive always present pectoral girdle.CHAPTER XXIII THE EVOLUTION OF MAN It has long been realized that the human body is constructed on the same general plan as that of other mammals. tion of man. 277 and 287). in most cases the feet as well. The mammae occupy a thoracic position. and the digits typically being furnished with nails. As a rule the young are born singly and are completely helpless during early feature. A The stomach is simple. with the carnivores and the horse with the fundamental structural resemblance. On the same basis that the dog is classified viz. only one pair being present. All but man. is perhaps the most outstanding feature of the primates. ungulates. the gorilla. plexity of the brain. the teeth In the lower monkeys 36 are also relatively unspecialized. monkeys. man must that order of be placed in the primate group mammals which The remarkable comincludes the lemurs. and apes. whose range has become considerably extended through artificial adaptation. teeth are present (Fig. that it comes into existence. 209). but although surpassing all the other mammals in mental development. 404 . which reaches its culmination in man. Distinctive Features of Primates.

by permission. the curvature of the spine. The general fundamental similarity the differences being mostly in details.) FIG. the position of the great toe. I . Among these. 272. of the teeth. note the size and shape of the cranium. "Organic Evolution" The Macmitlan Company. Vrom Lull. Skeletons of man and gorilla. the relative length rf the arms and legs. and of the lower jaw.THE EVOLUTION OF MAN 405 is striking.

marmosets (Hapalidae).406 FUNDAMENTALS OF BIOLOGY include six well-defined families: the lemurs The primates (Lemuridae). and mankind (Hominidae). 273. man Homo sapiens 1 variety. Mongolian. Old World monkeys (Cercopithecidae).) most closely related to man. The the lemurs comprise the lowest group. All the races of modern species are considered to belong to the same but three main varieties are commonly 1 recognized: the Ethiopian. In fact. the anthropoid apes the group FIG. the structural resemblances between the two highest families are much greater than between the anthropoid apes and the lower primates (Fig. South American monkeys (Cebidae). and Caucasian. The gibbon. Although the American Indian is sometimes separated as a fourth it is usually included under the Mongolian. Zoological Society. . the most primitive of the anthropoids in regard to certain characters of the skull and teeth but with limbs highly specialized for When on the gi ouiid it walks erect. (Courtesy of the New York arboreal life. the anthropoid apes (Simiidae). 272).

both in the temporary and Fio. rarely exceeding . the brain being more highly developed. the chimpanzee and gorilla in tropical Africa (Figs. (2) the number of teeth is the same as in the gibbon. and gorilla. are distinguished from the other primates by several important characters (1) All are tailless. The gibbon and orangutan are found in southern Asia. (3) the arms aro much longer than the legs. (4) the ground they can assume a semierect position on the hind limbs. Sumatra. 274. the 407 apes. (6) all are much more intelligent than the lower primates.) permanent sets. 273 to 276). New York Zoological Society. The gibbon is the smallest of the anthropoids. comprising The four anthropoid orangutan.THE EVOLUTION OF MAN The Anthropoid Apes. chimpanzee. : Old World monkeys and in man. (5) the thumb is short. but the great toe is well when on developed and opposable. a brown-haired anthropoid native of Borneo and The skull is higher and more rounded and the fare much longer than (Courtesy of the in the gibbon. The orangutan.

eye arches. Man. in the latter being developed as short tusks.) The skull of the gibbon is smooth on the top and does not have bony arches over the eyes. The arms are longer than the legs. In the chimpanzee and gorilla to maintain their balance. while the gorilla is the largest and most powerful. the male attaining an.408 FUNDAMENTALS OF BIOLOGY 3 feet in height. average height of about 5j^ feet and a weight of approximately 400 pounds. 277). and the eye arches are prominent (Fig. but relatively shorter than those of the gibbon and orangutan. Note the protruding jaw and veiy prominent. (Courtesy of the New York Zoological Society. Man is distinguished from the anthropoid apes and other primates chiefly by the following Distinctive Features of . but in the other anthropoids the skull bears a median crest. 275. The gibbon is the only form that habitually walks erect when on the ground. The chimpanzee. but the others can do so by touching the knuckles to the ground in order FKJ. the canine teeth are conspicuous.

apea. the lower jaw bears a distinct chin. (6) the arms are not so long as the legs. has four distinct curves. 276. and in Mounted specimen of mule goiilhi. adapted to erect posture.THE EVOLUTION OF MAN characters: (1) larger (3) 409 is The brain. (4) all the teeth are size. Chicago. the lai^esl of the anthropoid many ways the most highly specialized. but the great toe is not. especially the cerebrum.} the spinal column. (2) the face is short and nearly vertical. (7) the thumb is freely opposable. (Courtesy of the field Museum of Natural History. (5) FKJ. The differences between man chiefly associated with the and the anthropoid apes are assumption of the erect posture and . notably and more complex. (8) the power of articulate speech is fully developed. reduced in the canines being not larger than the others.

the largo canine teeth. Related to the development of the erect posture were such structural changes as the broadening of the pelvis. modified position of the head on the spinal column. it became possible for the precursors of man to make freer use of the hands. the carnivores and primates had a common In fact. the early employment of tools and weapons.410 FUNDAMENTALS OP BIOLOGY the development of the brain. the and the absence of a chin. etc. flat Note the very retreating forehead. For example. North in America and Europe during the Paleocene there existed a group of small arboreal mammals intermediate between the It will . but gradually enabling primitive man to communicate by means of language. History of the Primates. :uiiiV< ninir of the spine. marked the beginning of man's supremacy over the other mammals. origin. Associated with the development of the brain was the growing power of articulate speech. shorten- prominent eye arches. most likely in a primitive stock of insectivores. be recalled that the mammals began their ascendancy at the beginning of the Cenozoic era and soon gave rise to many diverse groups. the nose and protruding jaw. at first feebly manifested. According to prevalent scientific opinion. a most important factor in early human evolution. ing of the arms. correlated with increasing mental powers. With the gradual abandonment of arboreal life and the growing ability to walk erect with ease.

5. Prototypal anthropoid (Propliopithccus) of Oligocerie age. A. Ape man (Pithecanthropus) of Java.THE EVOLUTION OF MAN tree shrews (insectivores) 411 and lemurs (primates). Primitive primate principal branches of mankind and of the anthropoid apes. American Museum of Natural History. ancestor of the gibbons.} the primitive lemurs of Europe at that time. . (Courtesy of Professor William K. 278. Neanderthal man (Homo neandcrthalensis) 8. . Piltdown man (Eoanthropus) 6. . D. . 11. Gregory and C. Australian black man. Fig. representing the Negro group of races. one of the most primitive of existing human races. Heidelberg man (Homo heidelbergensis) 7. showing the evolution and relationships of the 1. American. Hottentot. Primitive anthropoid (Dryopithecus) from the Miocene of India. 3. Chinese. Family tree of man. representing the Caucasian group. 4. These forms have to the entire primate line. 12. 10. Cro-Magnon man (Homo sapiens) 9. 278. 2. One of them (Propliopithecus. ancestral been may The primates became differentiated as a distinct order of mammals during the Eocene. while the others are considered to have Remains of the . gorilla. gibbon. /?. In the Oligocene deposits of northern Egypt several different kinds of primates have been found intermediate between these primitive Eocene lemurs and the monkeys and apes of later times. many One kinds lived in North America and of these (Notharctus) is shown in Fig. chimpanzee. The fossil record shows that - fUjJV *#. FIG. 278) was probably a direct been forerunners of the Old World monkeys.*% mm. representing the Mongolian races. (Notharctus) from the Eocene of Wyoming. orangutan. from Egypt.

of about 1 million years duration. 279 and 280). them as ancestral to modern anthropoids but not to man. Pliocene. associated with the bones of extinct mammals. it and enormous continental consequence glaciers that of great climatic changes. they probably represent several divergent lines of descent (Fig. Asia. Our knowledge is based on fossils that have been found. has been called the "Age of Man. the remains of a creature that has been given the name of Pithecanthropus erectus (Figs. in river valley deposits or in caves. separated by intervening warm periods. he dominates the earth. Others think that one form (Dryopithecus. espeto which evidence points as the place ct' cially in central Asia stock caused a gradual disappearance of origin of the human has been suggested that this may have been the impelling cause of the origin of man. his immediate ancestors being forced to abandon arboreal existence and slowly to adopt a terrestrial mode of life in a progressively more open country. increasing aridity. and a femur (thigh bone). four distinct invasions of the ice occurred. The Quaternary period. it should not be assumed that they form a In fact. Fig. during the Miocene and early Pliocene. and Pleistocene formations of Europe. Substantiating this view is the fact that. 278) may represent a common Miocene stock from which both man and the higher apes arose. which they were found was at first supposed to be late Pliocene ." as it was during the Pleistocene that man began to rise in supremacy over the other mamThe mals. of course. and today.412 latter FUNDAMENTALS OF BIOLOGY group are represented in the Miocene. The first true anthropoids are recorded in the Miocene and Some authorities regard Pliocene rocks of Europe and India. These remains consist of a skull cap (roof of the skull) two upper molar The age of the deposit in teeth. Although we shall consider these races in the order of their probable age. it is generally agreed that single evolutionary series. Pleistocene epoch was characterized by the development of forests. and so it is customary to divide the Pleistocene of Europe into four glacial We are now ready and three interglacial stages. The Java Ape Man. In 1891 there was discovered in a river deposit in central Java. for the most part. and Africa. 278). to consider briefly some facts regarding tho earliest races of mankind. appeared at intervals as a In Europe.

Pithecanthropus erectus. of Java. retreating forehead and prominent eye arches. 279. as restored by ProAntiquity estimated at about 1 (Courtesy of Professor McGregor. the latter being almost as . while that of the lowest members of the human meters. Columbia University. The brain capacity has been estimated at 900 cubic centimeters. race (the native Australians) is about 1. fessor J.300 cubic centiThe average brain capacity of the white race is approxi- mately 1. The skull cap is very long in proportion to its breadth and is narrow in the frontal region.500 cubic centimeters and the maximum almost 2. the ape McGregor. H. possibly during the middle Pleistocene. authorities think that Pithecanthropus lived still later. Pithecanthropus had a very low. that of the male gorilla never exceeding 600 cubic centimeters.) man that of the chimpanzee. million years.THE EVOLUTION OF MAN Some 413 but was later established as very probably early Pleistocene.000 cubic centimeters. in the latter respect resembling FIG.

If . narrowness. by permission. "Men of the Old Stone Age. and the retreating forehead.the femur belonged to the same apes. From a study of plant and animal fossils later found in the same deposit to from which the Pithecanthropus remains were obtained. which we assume existed on the central Asiatic . Hence he was out of competition with his more progressive contemporaries. Judging from the slope of the forehead. it has been determined that the Java ape man was a forest-dwelling type. X J^. probably having migrated from a more northern region. (From Osborn. especially in the frontal region. greatly. 280. of modern man and the Side and top views of the skull cap of Pithecanthropus erectus. and so not in the direct line of descent. after Du Bois.) Fio." Charles Scribner's Sons. the creature was about 5 feet individual. its Note Most authorities regard Pithecanthropus as collaterally related modern man. tall 7 inches and walked erect. it is probable that the jaw protruded The teeth are intermediate in structure between thoec. the heavy eye arches. as seems probable.414 FUNDAMENTALS OF BIOLOGY conspicuous as those of the chimpanzee.

whose remains have been recently found in a cave deposit at Tien. The remains of more than 50 different types of mammals. pekinensia). The Peking Man. Elliot Smith in Scientific Monthly. including teeth and jaw fragments. China. being an almost complete uncrushed cranium (Fig. 280.) another primitive type similar in many respects to Pithecanthropus but generically distinct. near Peiping Chow Kow (formerly Peking). 281). all of which belong to . made in 1929.THE EVOLUTION OF MAN plains. 415 and remained primitive in an environment where livingwas easy. (From G. 281. is FIG. were associated with the Sinanthropus material. The remains consist of parts of 10 different skulls. besides other vertebrates. X H- Side and top views of the cranium of the Peking man (Sinanthropus Compare with Fig. The Peking man (Sinanthropus pekinensis). the most remarkable discovery.

England. is of a disour geologic time scale is correct. higher type.416 FUNDAMENTALS OF BIOLOGY Thus there is no doubt as to the very early Pleistocene. 282. as restored by Professor (Courtesy of Professor McGregor. . the Peking man resembles the next man to be considered.) J. The cranial capacity is approximately cubic centimeters. One of its most is the exceptional thickness of the cranium.000 fourths as high as in distinctive features modern man. H. Although the roof of the skull is slightly higher than that of Pithecanthropus. In form of skull and size of brain. while primitive. Sinanthropus shows a striking similarity to J'itluc<inthmpn}i. In this respect. but. of Sussex. The Piltdown man McGregor. as well as in the ape-like character of the jaw. The skull is narrow in front and provided with very prominent eye arches. it is only threetinctly 1. when the Peking man lived. and if remains are a million years old. these FIG.

berg man of (C). having been ascribed to the middle Pleis- tocene (third interglacial stage) by some.) The skull of the Piltdown man measures 11 to 12 millimeters in thickness. but the jaw to an extinct . that of the existing native Australians being 6 to 8 millimeters. 282). but the canines are very conspicuous. Heideland modern mafi (D). exceeded in thickness only by that of the The forehead is high and prominent and is withof is modern Europeans only 5 The cularly in the character of the lower jaw and the teeth. England. (From Woodward. Piltdown man (B).000 to 1. 283). young chimpanzee The teeth are human. out conspicuous eye arches (Fig. and In fact. The Piltdown man (Eoanthropus dawwas first known from a single broken skull and jawbone found in 1912 at Piltdown.300 cubic centimeters. Some authorities have considered the skull to belong to a primitive man. The primitive character of the Piltdown man is revealed partiformer to 6 millimeters. 283. found in association with the bones of extinct The remains were mammals and with some very primitive bone and flint implements. FIG. while recent evidence indicates that its age may be late Pliocene. the skull Peking man. is without a chin and is almost identical with that of a (Fig. Side view of jaws of chimpanzee (A).THE EVOLUTION OF MAN 417 The Piltdown Man. The age of the soni) material is uncertain. by permission. The cranial capacity has been variously estimated at 1. "Guide to the Fossil British Remains Man" Museum. to the early Pleistocene (first or second interglacial stage) by others. Sussex.

The Neanderthal Man. were it not for the teeth. after the original discovery. From the same formation have discovered in 1907 in southern FIG.) obtained the bones of many extinct mammals. "Men of the Old Schoetensack. after (From Osborn. The jaw was found in a river deposit 79 feet below the level of the and fourth glacial stages and is thought all . near Dusscldorf. particuThe Neanderthal race lived durlarly in southwestern France.418 FUNDAMENTALS OF BIOLOGY few years ape. a portion of another skull A and a tooth similar to that present in the first jaw were found in the same region. the jaw would have been regarded as that of an extinct anthropoid ape. by permission. was discovered in 1856 in the Neander Later many skeletons and Valley. The Heidelberg Man. claiming that their association was accidental. which are chin of modern man. bergensis) glacial The Heidelberg man (Homo hcidel- lived in period. The Heidelberg jaw is very large and heavy and. Germany. ing the third 'miergl:iri. The Neanderthal race (Homo neanderthalensis) is known from a number of specimens found mostly The first in caves in about 20 different localities in Europe. Age" Charles Scribncr's Sons. 284. lacks the projecting point that forms the In fact. a skull cap. more complete skulls were found elsewhere in Europe. however. specimen. as in the Peking and Piltdown jaws. so that there remains little doubt but that the skull and jaw belong to the same individual. Stone The Heidelberg jaw. Europe probably during the second interHe is known only from a single lower jaw Germany near Heidelberg (Figs. about X %. been distinctly human (albeit somewhat primitive in form). 283 and 284).

The eye arches were thick and very prominent. with a retreating forehead and a broad flat nose (Figs. the erect posture that characterized subsequent races not being fully established. (Courtesy of Professor McGregor. The Neanderthals were a low-statured people. The head protruded slightly forward from the broad rounded shoulders.THE EVOLUTION OF MAN to 419 perhaps have occupied Europe for at least 50.000 years ago. Head of the Neanderthal man.000 to 75. short and thick. the former being . modeled by Professor J.000 years twice as long. the males rarely exceeding 5 feet 5 inches in height. and narrow. The skull was thick. H. They walked with a stoop- ing gait. 285. McThis primitive race inhabited Europe 25.) FIG. The limbs were Gregor. 285 and 286). long. the knees being slightly bent forward as a result of the curvature of the thigh bones. The lower jaw and the teeth were primitive.

McGregor. the heavy eye arches. (Courtesy of Professor McGregor. Note the low forehead. with the missing parts (JR) restored by Professor J. ^ Fin.420 FUNDAMENTALS OF BIOLOGY Fro. France. 2b. . and the primitive jaw. Compare with Figs. X V^. H.) . X 277 and 286.Photograph ot a cast ol a Neanderthal skull found at La Chapellesux-Saints. in 1908. -Skull of modern man (Homo sapiens). Correze. 12S7. .

In development. shows that these primitive people knew the use Whether the Neanderthals became entirely extinct or partly evolved hit o the lower races of modern man is an unsettled question. any rate they were succeeded by a vastly superior type of people who came probably from Asia at the beginning of post-glacial approximately 25. but yet relatively and heavy. . the brain was essentially modern. This highly developed race. In some individuals the chin was slightly developed but in most ceases was absent. France.800 cubic The Cro-Magnons were a race of centimeters. being larger. Indeed. in fact.THE EVOLUTION OF MAN less 421 powerful than that of the Heidelberg man. being relatively smaller and the convolutions simpler. Neanderthal remai. but the canines were not so thick conspicuous as in the Piltdown man. with a broad face and high check bones (Fig. chiefly in caves.000 years ago. cattle. 288). intellectually they were vastly superior to the Neanderthals. the males averaging slightly over 6 feet in height. which is the seat of higher faculties. 1 :. The Neanderthals were a cave-dwelling race.. of fire. drawings. The skull was large but narrow. than that of the modern white race. but migrated into Europe and gradually replaced the Neanderthals. In some cases the cranial capacity was as much as 1. however. The lower jaw was heavy but was provided with a conspicuous chin. and although their stage of culture probably did not surpass that of some existing savages. and cave bears. were discovered at Cro-Magnon. A number of complete skeletons have been found time The Cro-Magnon Man.500 cubic centimeters. . of primitive flint tools They made u. ornamenting the walls of their caves with sculpture. belonging to Homo sapiens seems to have undergone its early evolution in Asia. of fire-charred bones of these animals with and weapons. They were tall in stature. in various parts of Europe.r wild horses. cave-dwelling hunters. The teeth were more primitive than in modern races. The proportions of the brain were different. The cranial capacity of the Neanderthals rarely exceeded 1. The first specimens These people were physically one of the most splendid races of man that has ever existed. and hunted such animals as The association reindeer.. The forehead was high and the eye" arches small or absent. this being the same as that of the average modern Caucasian. they were an artistic people. but ai. the frontal portion.

422 FUNDAMENTALS OF BIOLOGY and even with colored paintings. persisted in Europe for at least 10. H. Head of the Cro-Magnon type of Homo sapiens. almost entirely intellectual and social. Since the time of the Cro-Magrions. to human progress has been Lull says: . just as the Cro-Magnons themselves had done earlier. Restoration by Professor J.) Elements culture. however. bringing with them a higher FIG.000 years ago. McGregor. some extent they may have been absorbed by them. (Courtesy of Professor McGregor. They are thought to have Finally. of these new races are found among modern Europeans. 288. Although largely replaced by the races that succeeded them. they declined and became supplanted by four or five new races that came in probably from Asia. a race inhabiting southwestern Europe about 25.000 years.

THE EVOLUTION OF MAN Man's physical evolution has change is 423 any are: virtually ceased. His future evolution. and morally feeble. . . and of hand of sight. but in so far as being effected. and is thus contrary to the law of natural selection. must also in the long run have an adverse effect upon the race. in and progressive. skill. Such changes reduction of hair and teeth. smell. will be mental and spiritual rather than as such will be the logical conclusion of the marvellous results of organic evolution> . it by an enlightened eugenics is . unless offset so far as physical. largely for safety. mentally. it is largely retrogressive. and dulling of the senses and hearing upon which active creatures depend so That sort of charity which fosters the physically.


Cyaiiophyccac (blue-green algae) Class 2.) 4. PORIFKRA Sporozoa (Plasmodium. Schizomycetes (bacteria) Class 2. Ciliata (Paramecium. Angiospermae (aiigiosperms or flowering plants) Subclass I. (seed plants) Class 1. Anthozoa (corals and sea anemones) 425 . Monocotyledoneae (monocotyledons) THALLOPHYTA BRYOPHYTA SPERM ATOPHYTA SYNOPTIC CLASSIFICATION OF THE ANIMAL KINGDOM Phylum 1. Filicinae (ferns) Division IV. Fungi Class 1. Dicotyledoneae (dicotyledons) Subclass 2. Equisctinac (horsetails) Class 3. Algae Class 1. Lycopodinae (club mosses) Class 2. Rhodophyceae (red algae) Subdivision 2. Phy corny cetes (alga-like fungi) Class 4.) s Phylum COELENTERATA Class 1. PROTOZOA 1.APPENDIX SYNOPTIC CLASSIFICATION OF THE PLANT KINGDOM Division I. Mastigophora (Euglena and other flagellates) 3. etc. (moss plants) Class 1. II. Calcarea (calcareous sponges) Class 2. etc. Musci (mosses) Division III. etc. Class Class Class Class (unicellular animals) Sarcodiria (Amoeba. etc. PTERIDOPHYTA (fern plants) Class 1. Hydrozoa (Hydra and hydroids) Class 2. (thallus plants) Subdivision 1. Gymnospermae (gymnosperms) Class 2. Vorticella. Phylum (sponges) Class 1. Demospongia (fresh-water sponges. radiolaria. Phaeophyceae (brown algae) Class 4.) III. Myxomyeetes (slime molds) Class 3. Chlorophyceae (green algae) Class 3. Hexactinellida (glass sponges) Class 3. Scyphozoa (jelly fishes) Class 3.) 2. bath sponges. Basidiomy cetes (club fungi) Division II. fofaminifera. Ascoiny cetes (sac fungi) Class 5. Hepaticae (liverworts) Class 2.

etc. 1. Cestoda (tapeworms) Phylum VI. Provertebrata Subphylum 2. etc. ANNELIDA (segmented worms) Class 1. Class Class 2. 1.) 2. scallops) Gasteropoda (snails.) Class Class Class 1.) Cephalopoda (squids. mites. tunicates. ECHINODERMATA Asteroidea (starfishes) 2. Amphibia (amphibians) Reptilia (reptiles) Class Class Aves (birds) 5. Myriapoda (centipedes and millipedes) Arachnida (spiders. Class Class Class Ophiuroidea (brittle stars) Echinoidea (sea urchins and sand dollars) Holothuroidea (sea cucumbers) Crinoidea (sea lilies) Phylum XI. barnacles.) Pisces (fishes) 2. 5. Insecta (insects) etc. 3. MOLLUSCA Pelecypoda (mussels. Vertebra ta Class Class Class 1. Phylum XII. Hirudinea (leeches) Phylum X. Mammalia (mammals) . sandworms. clams. 4. ticks) 3.) 2. MOLLUSCOIDEA Bryozoa (moss animals) Brachiopoda (lamp shells) Phylum IX.) Class Class Class 2. CHORDATA Subphylum 1. 2. crabs. FUNDAMENTALS OF BIOLOGY CTENOPHORA (comb jellies) PLATYHELMINTHES (flatworms) Turbellaria (Planaria. V. Phylum XIII. 4. slugs. 3. etc. etc. Class Class Class Class Class Class Class 1. (lancelet. octopi. Trematoda (flukes) 3. nautili. scorpions.426 Phylum Phylum IV. oysters. TROCHELMINTHES (rotifers) Phylum VIII. Chaetopoda (earthworms. 3. 1. 4. NEMATHKLMINTHES (roundworms) Phylum VII. ARTHROPODA Crustacea (lobsters. etc.

225 Adrenalin. 312 Adaptive response. 225 Agave. animals. 89 significance. 137 Appendages. 316 fossorial. 182 circulation. 162. 101 acids. 407 Antibodies. 250 Andalusian poultry. 35 ancient. 29 Anaphase. crayfish. 157 Anther.INDEX (References in boldface type indicate pages on which illustrations appear. 49. earthworm. 38 simple multicellular. 231 Algae. 205 plants. 130 Annelids. 170 ventral. 153 fern. Java. 47. 390 inheritance. 86 Antheridium. 133. 311 volant. 25 locomotion. fern. 69 Annual ring. 170 Aortic arches. 309 natatorial. 24 Acorn-tongue worm. 84. 233. 170 origin. 36 Alimentary canal (see Enteron) Amoeboid c^ells. inheritance. 333 Anus. 70. 239 Antennae. 358. 290 arboreal. 32 groups. 238. 134. 48 Anthropoid apes. 22. 292 Albumen. 413. 151 Amphibians. 228 Angiosperms. 136 Annual plant. 139. 291 individual. 313 cursorial. 364 (see Amphioxus Lancelot) Amylopsin. 257 Alveoli. 166 Acquired characters. 54 moss. 208 Ambulacral groove. 249. 314 Addison's disease. 330 Ants and aphids. 149. 11 Amoeba. 36 vegetative body. 176. 312 Adaptations. spiny. 314 racial. 26. 165. 310 explanation of. 333 Alternation of generations. 131 Aorta. 389 Adaptation. 130. 41 sexual reproduction. 58 origin. 134 Ape man. 412. crayfish. 291 Adaptive radiation. 182 chief groups. 23 fission. 127 in seed plants. 130. 53 Anteater. 57.) Amino Absorption. 23. 329 Antigens. 311 convergent. 156. 225 Adrenal glands. 156. 204 Anal pore. 71 Annulus. 375 Annelida. 52. 172 dorsal. 414 Aphids. 37 colonial. 359 asexual reproduction. 121. 150 427 . 248.

332. 169 Beriberi. 69 Bile. grasshopper. 46 fore limb. 203 20 . Blood. 172. fern. 47. 216 Bread mold. 1 Appendix. 155. 66 Axon. 111. 209. 92 Archenteron (see Gastrocoele) Archeozoic era. of prey. 265. 149 general features. 205 duct. 357 Aristotle. 363 Arthropoda.428 FUNDAMENTALS OF BIOLOGY Biology. 164. 133. 123 Bat. vermiform. 209 corpuscles. 185 chief groups. 357 life. 121 yeast. 203 Arachnids. 63 Beetle. frog. 47 Budding. 114. 312 * man. 176. 342 Assimilation. 194. 146. 196 ancient. 170. 141 Bronchial tubes. 318 sporangium. 266. 185 dividing. 221 Auditory organs. storage root. 229 sponges. 123. 77 Blastocoele. 372 Archegoriium. 168. 43. metazoans. 146. 268 Bacteria. 162 Asparagus. 301 insect-eating. 125 Biennial plant. 200 sporophyte. 305 feet and bills. skull. 94 316 seedlings. flowers. 1 318 (see Box Acorn-tongue elder. 302 nectar-feeding. pod. 49 sexual organs. 177 man. 212 Blade. 168. 91. 168. 174. 161 Archaeopteryx. 91. 12. alternation of generations. 172 Axil. 235 Blastoporc. Blastula. 235 origin. 4 scope. 193. 361 Brain. 72 nitrogen-fixing. 302 swimming. 47 Beaver. 333 seed. 211 Bone. 49 vegetative body. leaf. 169 Arthrodirans. 157 Auricle. 304 Black-knot fungus. 60. 70. 234. root system with tubercles. 333 Brachiopods. 117. 44 Breeding. 280 Brittle star. air. 21 21* plasma. 310 Bean. 48 pine. 133. B Backcrossos. 171. 217 applications. 306 Beet. 359 nitrifying. 169 Bryophytes. Botany. leaf. 209 platelets. 305 wading. 14 Arteries. 64 moss. 99 coelenterates. 318. 165 Auditory canal. stem. 73 Basal disk. 303 371 302 scratching. plant and animal. 175 Kalanoglossus worm) Bark. 96. 323 Bladder. 371. 180 frog. 234.335 6. angiosperms. 146. 149 Arthropods. 206 Atrium. 196. 2 Birds.

71. 27 Ciliary muscles. cutting. 8 principle. 174. 251 man. . 235 cavity (see Blastocoele) Classification. 166 Cambium. 417 skull. 222. 8. 170. lateral. 171. 175. 5. 150 Carbohydrates. 90 Carrot. 7 wall. 70. 169 life. 159 Bulb. acquired. 361 339 341 Cleavage. 215. 297 metamorphosis. . 255. 377 54 Centrosomes. 250 reduction. 175 Cerebrum. 7 Chromosomes. 340. 11. 365 Carnivores. 259 map. 258 Chrysanthemum. 195 Castor bean. 18 C Cabbage. 174. 251 X. 248. 216 Cerebral hemispheres. 361 Choroid coat. crayfish. 150 Chlorophyll. 209 Circulatory system. 67 Cephalothorax. 250. 172 open system. 410 Chitin. 260 Cephalization. 62 Butterfly. 149. 223 Chromatin. 193. 133 grasshopper. 65 flower. 275 number. 6 earthworm. 408 jaw. 215. 6. 84. 301 head. 151 Capsule. 26. 216 Characters. 162 Cerebellum. teeth. vessels. 348 Cactus. root. 307 Carpel. 256 homologous. 5 body.and F-chromosomes. 106 Carboniferous and Permian life. 5 Centipede. 176. 157 mussel. 267. 254 generalized. 268 grouping in gametes. 128 Cephalopods. 223. 16 sap. 198 formation. 242 non-adaptive. 204 Cilia. winter. 169 Cellulose. wild and descendants. Bug. 293 Calyx. 2. 209 fishes. 74 Cell. 6 multiplication. 249. 150. 163 Chordates. 217 division. 98 Chyme. 170. 8 Chordata. cork. 49 Carapace. 224 Chimpanzee. 397 Chemical coordination. 234. 66 429 and branches. 170 closed system. 48.INDEX Buds. 266. 68 Buttercup. 73 63. 22. 194. 83 Chloroplasts. storage root. basis. 286 recessive. 170 mammals. 249 fruit fly. 364 Carboniferous vegetation. 63 Cartilage. 82. 72 Cambrian and Ordovician Capillaries. 67 67 dominant and 269. 145 vertebrates. 224 Circulation. 390 terminals. 88 66. Cenozoic era. 209 amphibians. 169 dead-leaf. 163 primitive.

100 Dermis. 223 Corolla. 96 Crab. 150 internal anatomy. 364 196 Diabetes. 131. 334 Cranial nerves. 90 Selaginella. 66 development of Clitellum. 73 Corn. 57 Cycle of elements. 313 Coral. 146. 132. 237 Collar cells. 149. Charles. in plants. 185. grain. 194. 163 hermit. Monterey. 5. 153 * Cone. 52 pine. 150 dissection. 222. 93. 193. 6 D Darwin. 236. 217 Dentine. 94 seedling. 131 Cloaca. 94. 132 formation. 33 formation. 151 Cretaceous life. 222 Coelenterata. 126 Decay Cork. 68. 122 Coelenterates. 391 Cytoplasm. 126 Color blindness. 152 external features. 395 Conduction. 120 Cotyledons.430 Climbing fig. 62. 76 Cornea. 422 Colony. blood. 223 Cultivation and domestication. 77 Cotoneaster. 292. 66 Conifers. 227 human. 79 Cuttings. 46 Cuticle. 167 Crossing over. 194 Connective tissue. 125 Coelom. 57 Conjugation. 279 Competition. 176. 216 Cranium. 61. Crop. 210 salamander. 195 Crystalline lens. 32 hydroid. 102 stem. 9 appendages. 32. 133. 225 Cro-Magnon man. 6 Cortex. 174. 70. 121 Colloids. 229 Conjunctiva. 274 Crow. frog. 133. Deliquescent branching. 349 Cup fungus. 124. 167. 320 Cyclostomes. 302 Crustaceans. 373 421. 209 frog. 317 Defectives. 287 Convergent adaptation. 177 sponge. 22. 104 278. 195 white fibrous. 69 Dendrites. 150 Cochlea. 111 Dietary requirements. 70. 221. 198 Diffusion. human. 83 Corpuscles. 60. 122. 181 Cypress. 168. 180. 213 Desert plants. inheritance. 123. Paramctium. 176. 96 stem. 169. 126 Coelenteron. restoration. 126 representative. branches. 73 . 176. 222 Connective and supporting tissues. 394 of organic matter. 202 Dependent plants. elimination of. 208 Diastase. 179 Crayfish. 101. 62 Diaphragm. 71. 194 yellow fibrous. 120. 98 Cycads. 293 Devonian fishes. 21. 130. Cretinism. FUNDAMENTALS OF BIOLOGY skeletonized leaf. lycopods and horsetails. 174. 71. root. 194 fatty. 73 cambium.

320 essential to plants. 324 Diseases'. 123. 369. teeth. 266. 329 Dodder. cycle. 201 extracellular. 131. 132. 201 intracellular. 238. 93 Dragonfly. 367. 201 Earthworm. 133 embryos. adaptive features. 130. 220 Egg. 133. 286 hereditary. 136 reproductive system. 23/ Diploid. 85. 75 Embryology and evolution. 181 Dinosaurs. stem. crayfish. 236 sponges. 93. 118. 135 * Echinodermata. 137. seeds. 235. 94 Drosophila (see Fruit fly) Duckbill. 386 Dormancy. 86. 118. 221. 311 Digits. 208 mussel. 323 infectious. 120 Endodermis. 124. Elements. 267. 134 nervous system. 202 Encystment. 384 stages in evolution. 230 plants. 125 coelenterates. 237 human. animals. 38 Ectoderm. 235. 141 representative. 157 man. 141 Ecology. 167 Dihyhrid. 308 Effector. 222 Eardrum. 170 Dominant and recessive characters. 167. cross section. 180. 128. 251 322 323 fungous. 23 . 346 Du jar din. 26 inner. 159 Embryo. 145 starfish. 133 129 168 grasshopper. 167. 62. 134 general features. 373. nephridium. 202. 127. 234. 17 Embryo sac. 123. 111 excretory system. 222 Earthworm. fern. 264 Elephant. 49 vertebrate. 250 human. 218 digestive system. 139 Digitigrade. 130 Digestive system. circulatory system. 63 Endoplasm. 87 Elasmobranchs. coelenterates. 48. 125 embryos. middle. 269. 133. 264 in protoplasm. 133. 133. 374 Diplohlastic. 88. 236 sponges. 221. 218. 222 222 Enamel. 3. 221.INDEX Digestion. 88. 203 plants. 133 dissection. 222 outer. 92 Dutchman's pipe. 110 Dogfish. 137 Echinoderms. 89. 239 Endocrine Ductless (see glands glands) 53 human. early. frog. tooth. 234. 371. 232. 124. Felix. 231. E Ear. 271 flat worm. 290 Ectocarpus. 233 later. 23 Edentates. 370. 240 moss. 385. 40. 132 diagram of nerve cord. bacterial. angiosperm. 111 431 animals. 345 Embryonic stages. root. 151 earthworm. 85. 114 Ectoplasm. 54. 132. Endoderm. 131 circulatory system. 269 ratio. 133 man.

100 conduction in plants.432 Endoskeleton. 166 human. 223 F Facets. 75 Fibrinogen. 26. 180. crayfish. 221. 337 Excretion. 231. 150. 169 man. 201 Epidermis. 110 storage. 228 Flagellates. 93. 31. impression. 403 early stages. 335 causative factors. 236. 199 Enzymes. 152 earthworm. 191 Equisetum (see Horsetail) Eras. 366 Feeble-mindedness. 152 earthworm. 287 Feeding habits. 30 Flagellum. 232 and cleavage. 170 Devonian. 73 Epiglottis. 354 Ferns. 100 . 109 reserve. 290 direct action. 338 organic and inorganic. 82. 337 nature of evidence. 62 formation. 150. 63. 317 Fern leaf. 222. 223 Eyelids. 164. 203 mussel. 205 stem. 151. 179 bony. 179. 83 Fluctuations. 41. 180 chief types. 51 Erepsin. 69 Exoskeleton. 263 external and internal. 134 frog. 303 Flower. 205 Esophagus. 400 Fly. 168 grasshopper. geologic. 55 Fertilization. 63 stem. 150. 72. 94 - Enteron. 172 grasshopper. 222 coats. 19. 111. 167 man. 158 gametophyte. 238 significance. 3. 167. 222. 133 frog. 50. 223 Eyeball. 83 parts. 137. 173 FUNDAMENTALS OF BIOLOGY 88. 390 organic and inorganic. 52. 212 mussel. 284 aims. leaf. 287 Euglena. 52. 11. 210 Filament. 146 Eugenics. 127 Flicker. 132. 208 Epithelium. 222 Evolution. 402 directive factors. 181 circulation. 109 Feces. 30. 211 Excretory system. 89. 79. 28. 345 evidence from classification. 31 Eustachian tube. 71. 6l. 180 Fishes. 222. 120 Flat worms. 251. 197 Fermentation. 52. 132 formation. 213 80 root. 133. compound. crayfish. 87. 54 spore-bearing structures. Endosperm. 156 Fats. 130. 190. 21. 379 importance. 237 Environment. 52 young sporophyte. 359 embryological evidence. 146 Excurrentjbranching. 121. 290 Eye. 364 Fission. 64 sexual organs. 30. 92. 336 popular misconceptions. 155 Food. 250 Fibers. 131. 339 horse and elephant. 84 Fins. 202. human.

339 Geographical distribution. cleavage. 271 Genus. 80 cells.INDEX Food. 176. 38. 263. 129 grasshopper. 167 heart. 277 Fungi. 133. 83. 168 skeletal system. 152 Galton. 41 Gamete. 172. 92 Ganglia. development transpiration. 264 combinations. 50 fern. 284 mussel. 235 Gastrovascular cavity tcron) (see Gastrula. 176 reproductive system. 173 Fruit. 24. 23. vith stomata. 44 Gastrocoele. 235 Gastric caeca. 166 blood corpuscles. 167. 136 Four o'clock. 49. 89 accessory. 267 chromosome map. 26. 181. primary. 44 vegetative body. 146 Ganoids. 45. length. 176 external features. 38 Gametophyte. fungous. 322. 146. 204 Gar circulatory system. 310 Fossils. 169 respirat ory system. 174. 364 pike. 44 saprophytic. 30 Foot. 229. 46. mollusks. 106 of cork. 90 Fruit fly. 266 Free assortment. 263. 48. 433 109 vacuoles. 121 Genes. 356 Geologic periods. 235 Gel. 194. 94 G Gall bladder. 196 brain. 165 207 323 crayfish. 152 earthworm. 237 Germination. 43 parasitic. 171 nervous system. 170 development. 90 aggregate. 145. 37. 28. 38 condition. 144. 319 spore production. 234. 35Ci Geotropism. 252. 167. 351 nature and formation. 406 Gill slits. inheritance. 276. 265 fruit fly. 177 respiratc ry organs. 48 seed plant. Francis. 349 Geologic eras. 147. 158 mussel. 246 Genotype. 178 digestive system. 52. 360 Geologic timf division. 234. 351 64 moss. 144. 49 Fore limbs. Gills. 148 Gametangium. 164. 175 flatworm. 168. 262. 262. 258 heritable variations. 276 chromosomes. 252. 235 Gastrulation. 263 Germ Germ layerp. later. 276 Genetics. 167 Gastric juice. 73 epidermis. 113 Geranium. principle. 351 types. 265 Frog. 46. 323 reproduction. 274 sex determination. 353 . 145 . 234. 89. utilization. 4 for. moss. 309. mammals. 88. Gibbon. 168 excretory system. 268 sex-linked inheritance. 253 primordial. 4. 203 Galls. 218 crayfish. 10 Gemmules. 90 true.

167 Glands. response. 146.434 Gills. 304 Green glands. 208 Glycogeri. 166 nervous system. 209 Hemaglobin. 16 Hookworm. H Habit. 291 Humus. 186 salivary. 251 Hawk. 123 reproduction. 172 mussel. 191 endocrine. 379 Eocene. 122 Hair follicle. 203 Heidelberg man. 167 mammal. 80 Hooke. 192 Glochidium. 67 Hinge ligament. 151. 170 longitudinal section. 51. 245 and environment. 272. 90 78 Horsetails. 233 hind leg. 158 respiratory system. 264 Honeybee. 99 Grass seedling. 67 Habitat. 213 types. 157 dissection. 55. nest. 285 Hereditary elements. 147 Glottis. 75 Herbaceous perennial. Robert. 273 Gymnosperms. 133. 383 Guinea pig. 143 Homospory. 158 external features. 228 Haploid. 132. 171 grasshopper. 298 respiratory system. 113. 114 Gray matter. 381 stages in evolution. 126. 146 Heartwood. crayfish. 214 sweat. 129 Hormones. 166 mouth parts. 247 Hereditary characters. 169. 283 Hydra. 206 Goiter. 178 Gizzard. 146. 69 Grasshopper. 133. 209 sebaceous. 307. 60. 223 vitreous. 146 circulatory system. 366 Host. 126 . 69. 56 Heterozygous. 69 Hereditary bridge. 228 Guard cells. Herb. 167 excretory system. 220 Grebe. 123 Hydroid. frog. 57. 262 Hybridization. 223 leaf. Horse chestnut. 406. 225 Gorilla. . 309 skull. 158 reproductive system. starfish. 68 primitive. 135 Hetcrospory. 158 Gravity. 157 digestive system. 366 reproductive features. 264 Hickory twig. 224 Horse. 418 Hemacyanin. aqueous. 214 fission. 224 Heart. 125 structure. 152 Grouse. 215. 303 Heart. 162 fish. 2. 417. 264 Heredity. adaptive features. 60. 303 Humor. 213. 285 Hermaphroditic. 381 fore limb. 101 Hybrid. root tip. 243 Growth. FUNDAMENTALS OF BIOLOGY 139 tadpole. in man. 213. 322 Hummingbird. 55 Homozygous. 222 mammary. inheritance. 207 409 Grafting. 380. 73 lachrymal.

389 Andalusian poultry. 262. 172. 241 Larynx. 143 Iris. 100 Labium. 319 Indusium. 79. 133 flatworm. 132. frog. 82 L 21. 223 Light. frog. 86. 92 Intercellular spaces. 329 Independent plants. 116. 64. 192 Kidneys. to gravity. 300 Keratin. 263. 117 Hypocotyl. 116) 116 skeletonized. 151. 140. 162 arrangement. 86 pollen grain. 332 Life. 285 pea. 278. 226 Integument. 203 mussel. 167. 242 Larval stages. 147. 77 internal structure. anther. 86. 176* 177 man. 347 Internode. 97 435 K Katydid. 272. 65 cross section. 267. 146. 246 Insects. 246 color blindness. 152 representative. 96 arch. 12 223 Irritability. 67. 270 sex-linked. 146 Hypostome. 168 grasshopper. 93. 77 Leaflets. 124 I Ichthyosaurs. 331 cross section. 368 Immunity. crayfish. 215 126 Jiirassic life. acquired characters. 2. Carl 267 von. 206 Inheritance. Leech. 52 Inflorescence. 169 Insulin. 79 earthworm. 263 biparcntal. 81 Lamarck. response. 212 mussel. 278 uniparental. 73 Lichens. 79 divided. 167 man. 146 starfish. 313. 77 functions. nature of. Jean Baptiste Lamprey. 277. 390 young embryos. 266. 136 Lenticels. 116 Lily. 276. 181 Lancelot. 166 Labrum. 274 Linne". 167. Ligament. 174 Linkage. 205 Lacteal. 3 . suspensory. 240 Invertebrates. 273 man. 65 Intestinal juice.INDEX Hydrotropism. 236 de. 164 older embryos. 87 Limbs. 12. 169. 208 Lateral line. 166 Lactase. 78 epidermis. Indian pipe. 367. 139 Intrauterine development. 80 external features. 114 to light. 279 four o'clock. 266 guinea pig. 113. 94. 77 Intergrading species. 79 response. 205. 179 Leaf. 128 frog. 136. 234 Larva. 202. 86 ovule. 205 Intestine.

Liverworts. r limbs. 327 Malpighian tubes. 210 Man. 202 Lung books. 174. 324. 85. 281 Mastodon. 285 intestinal villus. 13 Lymphatics. 411 hereditary characters. 262. 100. 248. 30 Malarial parasite. 364 Mistletoe. 366 Metabolism. 261 Mendelian principles. 263 Maxillae. 182. 169. 119 fission. 228 Miastor. 326 Mitosis. 144 Marchantia. 202. 28. 160. 46 Marsupials. 367 placental. 156 Mechanistic theory. 206 Metamere. 126. 216 chromosomes. 237 Mcsoglca. blood brain. 26. 86. skeleton. 197 Metagenesis. 326. 249 Mohl. 223 family tree. 46 Millipede. 270 assort- segregation. 198 digestion. 240 skin. 167. 238. 212 eye. 236 formation. 184 420 tooth. 239 Mass culture. 263. 186 archaic. 187 Membranous labyrinth. 213 teeth. 305 Mesoderm. 384 corpuscles. mucous. 364 Lungs. 216 175. 30 Micropyle. 222. 26. 168. 239 generalized. 92 Microspore. 387 Maturation. 46 Lizard. 260 Metazoans. 221.436 Liver. 127 Mammary Mammoth. 269. 187. 417 nervous system. 346 excretory system. 308. 144. 203 digestive system. glands. 2. 89 Membranes. 89 Mildew. 17 Moisture. 117 jaw. 310 origin. germ cells. ment. 211 hearts. 215 . 127 Megaspore. 222 free egg-laying. 203 distinctive features. 191 Maltase. 158 Medulla obloiigata. 305 Mendel. 248. 215. 128. 186 362. 378 chief groups. 252. 131 Metamorphosis. 190 serous. 26. 259 dietary requirements. 120. Grcgor. 211 Mandibles. 252 Micronucleus. Hugo von. 205 Mammals. impression. 230 embryo. 211 spaces. 124 Mcsophyll. 168. 27. 211 M Macroruicleus. 60. Medusa. 56. 123. 242 Metaphase. 243 butterfly. 207 Lycopods. 408 ear. 309. response. 101 Lungfishes. 146 cavity. 240. 366 Lymph. 79 Mesozoic era. 51. 88. 154 Miocene fish. 203 FUNDAMENTALS OF BIOLOGY Man. 211 nodes. 56. 181. embryo. 166 Mantle. 249. 406 skull. 222 egg and sperm.

176. 166 formation. 236. 220 starfish. 165 starfish. 146 reproduction 146 arid earthworm. 218. 215 crayfish. 193 Mud 139 central. 133. 144. Nervous system. 164. 375. 310 fore limb. 219 motor. 152 Mushroom. 144 larva. 163. 147 representative. 223 sensory. 6 Mosses. 308 skull. 129 grasshopper. 237 peripheral. 136 flatworm. 400 Mutations. flower. 158 lancelet. 134 Nerve. 147 mantle cavity. 46 Mussel. 376 Moss. Morphology. 143 Mollusks. 133. 218 flatworm. 420 Nemathelminthes. 5 Nuclear sap. 93 stem. 400 Node. 176. 132. 164 Myxedema. 215 development. 162 earthworm. 297 Natural selection. 144 nervous system. 397 and mutation. 344 N Nasturtium. leaf cells. 187. 7 Nucleolus. 167. grouse. Mouth 130. 158 man. 194. 401 theory. 146 Muscle tissue. 193. 217. 194. 238 fibers. 437 352 Mole. 2 Mosasaurs. 218 auditory. 143. 400 objections. 128. 399 Mutation. 269 Monotremcs. 393. 76 Monohybrid. 124 Nematode worm. 343. 319 Myotomes. 165. 243 Neuron. 65 North America in Cambrian time. Neanderthal man. 76. 48 mixed. 398 and natural selection. 221 cell (see 306 Neuron) 218 Monocotyledons. 176. 136. 318. 143 fresh-water. 148 Monkey. 237 Nuclear membrane. dissection. 218 motor. 215 mussel. 176. 220 Nictitating membrane. 46 life history. fore limb.INDEX Molds and casts. 165. 176 grasshopper. 164. 217 intermediate. 133. 366 Notochord. 282. 164. 218. 396 limitations. crayfish. 131 parts. Mutant. 419. 44. 194. 146 origin in chordates. 130 Nephridium. 176. 310 Mollusca. 176. 143. 183 Muscle. 7 . Mouth. grasshopper. 192. 129 frog. 418. 193 Nerve cord. 179 Myriapods. natural. 146 external features. 166 puppy. adductor. 400 Mycelium. 43. 218 olfactory. sensory. 176 optic. 217 Nest. 218 tissue. 225 139 sympathetic. 129 Nematocysts.

155 Octopus. 120 Osmosis. 173. 270 Pectoral girdle. 83. 322 malarial. 238 Ovipositor. 157 male. 126 Ocelli. 26. 174. 27 Pelvic girdle. 91. 133. 351 Paleozoic era. 5. 30. angiosperm. 173. 123. 88 tube. 351. 167. 345 Oocytes. 267. Parenchyma. 1 multicellular. 204 Oedogonium. 167. 204 Perch. 69 Perianth. 27 conjugation. 103 Ostracoderms. angiosperm. 102. 175 Pedigree culture. 80 spongy. 360 Pancreas. 169. 28. 86. locomotion. 28 Orange. 80 Parental care. 361. 266. 263. 360 . actual. 255 Oogenesis. 28. 27 Parasite. 140 Overproduction. 282 Peking man. 84 Pericardium. 125 mussel. 174. 71 72 Orangutan. 243 Parthenogenesis. 336. 68 Paramedum. 14. 62. 407 Organic remains. 88 crayfish. 133. 86. 57. 169 Periods. bulb. 7. 167. 148 (see Egg) Oxyhemaglobin. 180 Perennial plant. 8 vegetative. 394 Overspecialization. 135 frog. 201 Pellicle. 83 cortical. 3^8 Pepsin. 210 Ovum Paleontology. 92 O Oaks. 31 Oral groove. inheritance. 397 Parthenogenetic animals. 146 starfish. 352 Organisms. 90. 333 Parathyroid glands. 153 earthworm. 160 Oviparity. 40 Oenothera (see Primrose. 177 grasshopper. 150 Ovule. 252. 340 Obdia. 58 Osculum.FUNDAMENTALS OF BIOLOGY Nucleus. 18 palisade. 26. 175 Pentadactyl. earthworm. 203 Pancreatic fluid. 160 hydra. 168. 87 Nymph. 324 Parasitic plants. 34 unicellular. 225 Optic lobes. 87 Oviducts. 232 Organs. 84. 100. 15 Peas. 145. 83. 327 Parasitic animals. Louis. 79. 175 Onion. 87 pine. 233 Pasteur. six species. 263 Operculum. 175 Oral funnel. 87 primary endosperm. 26. 135 frog. 79. flower and fruit. 6 generative. 253 Oogonia. 362 Ovary. 178. leaf. 139. evening) Oleander. fission. 321 reciprocal. 177 grasshopper. later geologic. 294 Olfactory lobes. 145. 26. 180 root-tip cells. 322 Parasitism. 416 Pellagra. 229 29 248 Ontogeny. 252. 84. 326.

345 Physiology. 366 Peristalsis. 81 root. 226 Placenta. 87 . 104 Platyhelminthes. 19. 51 Plasma. 303 insect. 439 Plesiosaurs.. 12 structure. 87. 10 motion. 109 Proterozoic era. 358 Protococcus. 83. 416. 48. 90. 52. 92 cross. 305 Plants and animals. 133. 55 spore production. 84 Petiole. tuber. 128 fission. 63 stem. 367. 87 Pollen. 96. 86. 70. 107 Porpoise. 23. 249 Prostomium. 91 ovule. 86. 5. Java) Pituitary gland. 305. 19 Protonema. 410 306 109 Primrose. 108 process. 297 Prophase. 79 and respiration. evening. 51 Plasmolysis. 402 history. 352 Petrified logs. 164 Pseudopodium. 119 Provertebrata. 198 formation. 146. 106 release of oxygen. 77 Plumule. 8 Protozoa. 357 life. 353 Petunia. 72 Photosynthesis. 399 Privet. Petrifactions. 131. 48. 241 Planaria. 132. 10. raw materials. 84 compound. 76 Pithecanthropus (see Ape man. 52 vegetative body. 52 heterospory. 255 Polar caps. 336. flower. 2 among insects. 11 composition. 93. 766 Plexuses. 94. 61. 77. 84. 98 Primates. 68 cones and cone scales. 84 Pith. 248. 133. 6. 85 86 296 self. 127. 2. 107 tube. 299 Piltdown man. 113 Phototropism. 106 agent. 313 Potato. 133 Protective resemblance. 228 Plantigrade. 209 Plasma membrane. 18. teeth. 52 leaf. 319 Pistil. 217 Petal. 108 necessity of light. 96 Polar bodies. 300 Proteins. Porifera. 375. 23 Plasmodium (see Malarial parasite) gametophyte. 263. 126 Poorwill. 107 equation. 8. 71. 298 grain. basket. 329 Pharynx. 115 Phylogeny. 70. 72. blood. 163 Provertebrates. 164 Phenotype. leaf. 118 resemblances. 5. butterfly. distinctive features. 83. skeleton. 79. 3 Proboscis.INDEX 204 Permian amphibian. 85 Polyp. 127 . Pollination. 240. 62. 24 Pteridophytes. 84 simple. 1 19 energy. 88. 249 Polar nuclei. 271 Phloem. 417 Pine. 49 Protoplasm. 71. differences. 92 Pinesap. 298 Phagocytes. 17 behavior. 128. 68.

208 internal. eyeball. 126. 343 Rodents. 375 Ptyalin. leaf infected with fungus. 203 stem tip. 203 cells. 206 external. Schleiden. 43. 153 earthworm. 157 sexual. 141 Sand worm. 117 to moisture. 100. 161 Scurvy. Quaternary 379. 317 Pyloric caeca. 73 Schultze. 2 asexual. 319 Saprophytism. 218 arc. 344 chromosomes. 16 Max. 367. 78 Rostrum. 204 Reduction of Sage. 83 system. 111. grasshopper. 16 Sclerotic coat. 251. 141 . 61 Recapitulation principle. 140 Reptiles. 21. 58 hairs. 58 tip. 176 grasshopper. 37 in animals. 16 Schwann. 184 origin. 135 frog. 222. 365 Respiration. teeth. 51. 60. 242 Pupil. 38 in animals. 317 Sap wood. 59 functions. blood epithelium. 223 Putrefaction. 66 Sand dollar. 12. 158 mussel. 334 Sea cucumber. 223 Scorpion. 223 Rhizoid. 149. 319 Saprophytic seed plants. 117 Quail. Roundworms. 97. 146 starfish. 114 to light. 47. 299 Salamander. 412 R Radish. 264 Reflex action. 58 cross section. 69 Rootcap. Receptacle. to gravity. 6 Salamanders. 61. 54 Rhizome. 14 man and gorilla. in algae. Receptor. 204 Pulp cavity. Francesco. 150 Rose. 62 fibrous. 303 storage. 53. 222. 306 Root. 183 Saliva. 191 skin cells. crayfish. vestigial hind limbs. 63 life. 98 345 S Sacrum. 6 Reproduction. 204 Python. 219 Rennin. 129 Runner. 68 man. 136 Salvia.440 FUNDAMENTALS OF BIOLOGY Rhizopus (see Pterosaurs. 95. strawberry. tooth. 218. 69. 322 seedlings. 200 Sea anemone. 228 Salivary glands. 116. 208 Retina. 69 leaf. 220 Redi. 202 Bread mold) Rickets. 183. in algae. 200 Pupa. 229 Reproductive system. 173 Saprophytic fungi. Theodor. 184 chief groups. Matthias. flower. 374. 62 response. 113. Saprophytes. 139 Pyloric valve. 61 mature.

340 equilibrium. 393. 50 dependent and independent. 299 Sol. 263. 58 Sepal. 57 saprophytic. 145 Skeleton. 49. 68 Somatic cells. 263. 174 Slug. 296 formation. 84 Spermatozoan Spider. 252. 165. 277. moss. 174. 144. 120 cross section. appendicular. 120 Sinuses. man and 405 Spontaneous generation. 231 fern. 48 seed plant. 158. 53 Skin. 121 life. 181. 176. 55 Smut. 161 (see Sperm) Serum. 396 Semicircular canals.INDEX Sea squirts. 252 Somite (see Metamere) Sori. 166. 95 441 342 Snapdragon. 136 frog. 395 intergrading. 95 Sperm 130. 207 Spirogyra. 49 Spicules. 57. 42 Secretin. 157. 74 Silurian Spleen. 222 Sense organs. rhizome. cumulative effect. 311 fore limb. 160 66 Spermary 398 Selection. 152 Spongin. frog. 92 structure. 295. 313 Shoot. 37. 132 Spinal cord. 88 gymnosperm. 255 Spermatogenesis. 221 Spermatocytes. 94 Seed ferns. 364. 93 dispersal. 221. 213 cross section. frog. 40 Snail. 258 Sex-linked inheritance. 230 moss. 176. 43. 89 young fern. 119. 178. 217. 52. 55 fern. 46 Sport (see Mutant) . 339. 179. 38. 226 Seed. 213 Skull. ducts. 176. 131. 347 Sperm. 174 axial. 52. 211 Seta. 278 Shark. 220 Spinal nerves. 137. 253 Spermatophytes. 139 Sieve tubes. 265 grasshopper. 141 Seal. 176. 310 Seaweeds. 53 Species. 83. 146. 91. 276. 319 Seedling. alga. 64 system. 252. 96 corn. natural. corn. 249 Spiracles. 88 bean. 10 Solomon's seal. animals. 93. 121 Setae. 148. 53 Spore. 46. 89. 120. 253 Spermatogonia. bipolar. 48. 173. 220 Spindle. 216. 38 Sporophyte. 121 Siphon. 230. 135 Segmented worms. 176 (see Testis) Segregation. 120 reproduction. 48 seed plant. 39 Sex determination. 14 Sporangium. development. and Devonian 363 structure. 305 Sieve plate. Selaginella. earthworm. 342 moss. 174 173 gorilla. 44. 168 Sponges. 365 Seed plants. earthworm. 54 human. principle. 262. 166 Sea urchin. 64 Shrew.

158 hydra. 224 Tissues. 70. 129 Trachea. 304 Teeth. 204 Stem. FUNDAMENTALS OF BIOLOGY 6 Stamen. to gravity. 187 cell. 68 Sternum. 8. 328 Taproot. 213 Sweet pea. 248. 123 Tertiary life. 207 Tracheids. 168 grasshopper. 158. 72. 151 Tentacles. 167 84 Sucrase. 190 animal. 181 Thallophytes. 221 Taxonomy. 225 Thyroid gland. 140 Stomach. 83. Superiors. 72. increase of. 135 frog. 8. 3 Teal. 190 conducting. 84 Stilt. 114 to light. 139 btomata. 202 Teleosts. 145 starfish. 29 . 167. 75 monocotyledons. 140 structure. 83. 72 earthworm. 84. bilateral. hydra. 120. 253 Systems. 202. 364 Telophase. 202 Tortoise. 65 underground. 6 Tadpole. 75 muscle. 88. 211 Thymus gland. 146 starfish. 59 Taste buds. 74 Transpiration. 330 Symmetry. 304 Stipules. 71. 64 functions. 193 Tongue. tip. 78 70. 153 reproduction. 79. 64 response. human. 162. 128. 139 egg. 182. 33 Thoracic duct. 149. crayfish. 289 Survival of the fittest. 176 grasshopper. 299 Symbiosis. 97. 69 herbs and woody vines. 140 Steapsin. 206. 137. 129 Trichocysts.442 Squash. 174 Stigma. 184 Sweat glands. 131 radial. 83. 193 nerve. rudimentary. 169 Tracheae. 98 Struggle for existence. 249. 203 mussel. 93. 123 Synapse. 26. 377 Testa. 151. crayfish. 138 dissection. 116 structure. 194 mechanical. 129 Trichinosis. 178 Tail bones. 137 water. 76 young woody. 396 Tooth. 77. 218 Synapsis. 105 Triassic life. 73 connective and supporting. hair Squirrel. 167 man.vascular system. 395 Sturgeon. 125 mussel. insects. 344 Tapeworms. 35 Thallus. runner. flower. 250 Telson. 224 Thyroxin. 367 Trichinella. 94 Testis. 192. 90 Starfish. 116. 152 frog. 104. 80 Stone cells. 123. 157 man. 75 Strawberry. frog. 205 Style.

and non-heritable. 330 Vacuoles. 169 leaves. 26. 281 81 root. 173. 146. 400 fruit fly. 52. 362 443 Veins. 20 'Yolk. 68. 299 329 U Ulothrix. 179 Vessels. 238 Vorticella. 79 Vascular cylinder. 24. " 30 398 W Walking-stick insect. 392 Uterus. 143 Virus. 280. 68 Tunicates (see Sea squirts) Twins. 71. 136. 140 Trypsin. stem. 215. 6. flower. 342 Villus. 143. 240 Vries. 140 Tuber. Ungulates. 133 Vertebrae. 398 heritable origin. 163 groups. frog. teeth. 79. 212 Urine. 71. 361. 309 skull. 174 Use and disuse. 231. 52 leaf. 172 Triploblastic. 1 Zygote. 28. 174. 173. human. pulmonary. 76. 22. 212 mussel. root. 23. 68 Trilobites. 300 Water mold. 200 Xylem. 29. food. Hugo de. sporangium. 61. animals. 163. 23. leaf. 241 Urabo. 31 (see Xylem) X Xerophthalmia. 146. 140 White matter. 132. 72. 70. 36. 128. 37 Umbilical cord. 30. 163. 7 contractile. 398 Yeast. 308 Unguligrade. 170. 72 274 germinal and somatic. 13 Vitamins. 44 Water-vascular system. 78. 25. 75 Violet. 61. 172. 311 Urea. 204 Tube feet. xylem. 62 stem. 171 systemic. Vitalistic theory. 28.INDEX Trihybrid. 154. 212 Ureters. 63 continuous and discontinuous. 72 Vegetative propagation. 240. 62. 76 Vascular rays. 260 Typhlosole. 237 Trochophore. 79 Zoology. 176 man. 77. 212 Urostyle. 207 Vines. 307 Wood Vaccination. 26. 220 Willow. 77 Wolf. 171 Ventricle. 21 . 146 Urethra. 262 397 Vascular bundles. 97 Veins. 179 Vertebrates. 74 Vestigial structures. 395 Variations. 137. ^0 Variation. 71. 199 Viviparity. 206. 174 Vertebral column. 171. filterable. 236. 177. 70. 39 Zymase. 76. 272 Trillium. fore limb.