morphology

—morphologic /mawr'feuh loj"ik/, morphological, adj. —morphologically, adv. —morphologist, n.

/mawr fol"euh jee/, n.

1. the branch of biology dealing with the form and structure of organisms.

2. the form and structure of an organism considered as a whole.

3. Ling.

a. the patterns of word formation in a particular language, including inflection, derivation, and composition.

b. the study and description of such patterns.

c. the study of the behavior and combination of morphemes.

4. Physical Geog. geomorphology.

5. the form or structure of anything: to gain an insight into the morphology of our political system.

6. the study of the form or structure of anything.

[1820-30; MORPHO- + -LOGY; first formed in G]

* * *

I

In biology, the study of the size, shape, and structure of organisms in relation to some principle or generalization.

Whereas anatomy describes the structure of organisms, morphology explains the shapes and arrangement of parts of organisms in terms of such general principles as evolutionary relations, function, and development.

II

In linguistics, the internal construction system of words and its study.

Languages vary widely in the number of morphemes a word can have. English has many words with multiple morphemes (e.g., replacement is composed of re-, place, and -ment). Many American Indian languages have a highly complex morphology; other languages, such as Chinese, have a simple one. Morphology includes the grammatical processes of inflection, marking categories like person, tense, and case (e.g., the -s in jumps marks the third-person singular in the present tense), and derivation, the formation of new words from existing words (e.g., acceptable from accept).

* * *

▪ biology

Introduction

 in biology, the study of the size, shape, and structure of animals (zoology), plants (botany), and microorganisms and of the relationships of the parts comprising them. The term refers to the general aspects of biological form and arrangement of the parts of a plant or an animal. The term anatomy also refers to the study of biological structure but usually suggests study of the details of either gross or microscopic structure. In practice, however, the two terms are used almost synonymously.

      Typically, morphology is contrasted with physiology, which deals with studies of the functions of organisms and their parts; function and structure are so closely interrelated, however, that their separation is somewhat artificial. Morphologists were originally concerned with the bones, muscles, blood vessels, and nerves comprising the bodies of animals and the roots, stems, leaves, and flower parts comprising the bodies of higher plants. The development of the light microscope made possible the examination of some structural details of individual tissues and single cells; the development of the electron microscope and of methods for preparing ultrathin sections of tissues created an entirely new aspect of morphology—that involving the detailed structure of cells. Electron microscopy has gradually revealed the amazing complexity of the many structures comprising the cells of plants and animals. Other physical techniques have permitted biologists to investigate the morphology of complex molecules such as hemoglobin, the gas-carrying protein of blood, and deoxyribonucleic acid (DNA), of which most genes are composed. Thus, morphology encompasses the study of biological structures over a tremendous range of sizes, from the macroscopic to the molecular.

      A thorough knowledge of structure (morphology) is of fundamental importance to the physician, to the veterinarian, and to the plant pathologist, all of whom are concerned with the kinds and causes of the structural changes that result from specific diseases.

Historical background

      Evidence that prehistoric man appreciated the form and structure of his contemporary animals has survived in the form of paintings on the walls of caves in France, Spain, and elsewhere. During the early civilizations of China, Egypt, and the Near East, as man learned to domesticate certain animals and to cultivate many fruits and grains, he also acquired knowledge about the structures of various plants and animals.

       Aristotle was interested in biological form and structure, and his Historia animalium contains excellent descriptions, clearly recognizable in extant species, of the animals of Greece and Asia Minor. He was also interested in developmental morphology and studied the development of chicks before hatching and the breeding methods of sharks and bees. Galen (Galen Of Pergamum) was among the first to dissect animals and to make careful records of his observations of internal structures. His descriptions of the human body, though they remained the unquestioned authority for more than 1,000 years, contained some remarkable errors, for they were based on dissections of pigs and monkeys rather than of humans.

      Although it is difficult to pinpoint the emergence of modern morphology as a science, one of the early landmarks was the publication in 1543 of De humani corporis fabrica by Andreas Vesalius (Vesalius, Andreas), whose careful dissections of human bodies and accurate drawings of his observations revealed many of the inaccuracies in Galen's earlier descriptions of the human body.

      In 1661 an Italian physiologist, Marcello Malpighi (Malpighi, Marcello), the founder of microscopic anatomy, demonstrated the presence of the small blood vessels called capillaries, which connect arteries and veins. The existence of capillaries had been postulated 30 years earlier by the English physician William Harvey (Harvey, William), whose classic experiments on the direction of blood flow in arteries and veins indicated that minute connections must exist between them. Between 1668 and 1680, the Dutch microscopist Antonie van Leeuwenhoek (Leeuwenhoek, Antonie van) used the recently invented microscope to describe red blood cells, human sperm cells, bacteria, protozoans, and various other structures.

      Cellular components—the nucleus and nucleolus of plant cells and the chromosomes within the nucleus—and the complex sequence of nuclear events (mitosis) that occur during cell division were described by various scientists throughout the 19th century. Organographie der Pflanzen (1898–1901; Organography of Plants, 1900–05), the great work of a German botanist, Karl von Goebel (Goebel, Karl von), who was associated with morphology in all its aspects, remains a classic in the field. The Scot John Hunter (Hunter, John) and the Frenchman Georges Cuvier (Cuvier, Georges, Baron) were early 19th-century pioneers in the study of similar structures in different animals—i.e., comparative morphology. Cuvier in particular was among the first to study the structures of both fossils and living organisms and is credited with founding the science of paleontology. A British biologist, Sir Richard Owen (Owen, Sir Richard), developed two concepts of basic importance in comparative morphology—homology, which refers to intrinsic structural similarity, and analogy, which refers to superficial functional similarity. Although the concepts antedate the Darwinian view of evolution, the anatomical data on which they were based became, largely as a result of the work of the German comparative anatomist Carl Gegenbaur, important evidence in favour of evolutionary change, despite Owen's steady unwillingness to accept the view of diversification of life from a common origin.

      One of the major thrusts in contemporary morphology has been the elucidation of the molecular basis of cellular structure. Techniques such as electron microscopy have revealed the complex details of cell structure, provided a basis for relating structural details to the particular functions of the cell, and shown that certain cellular components occur in a variety of tissues. Studies of the smallest components of cells have clarified the structural basis not only for the contraction of muscle cells but also for the motility of the tail of the sperm cell and the hairlike projections (cilia and flagella) found on protozoans and other cells. Studies involving the structural details of plant cells, although begun somewhat later than those concerned with animal cells, have revealed fascinating facts about such important structures as the chloroplasts, which contain chlorophyll that functions in photosynthesis. Attention has also been focussed on the plant tissues composed of cells that retain their power to divide (meristems), particularly at the tips of stems, and their relationship with the new parts to which they give rise. The structural details of bacteria and blue-green algae, which are similar to each other in many respects but markedly different from both higher plants and animals, have been studied in an attempt to determine their origin.

      Morphology continues to be of importance in taxonomy because morphological features characteristic of a particular species are used to identify it. As biologists have begun to devote more attention to ecology, the identification of plant and animal species present in an area and perhaps changing in numbers in response to environmental changes has become increasingly significant.

Fundamental concepts

homology and analogy

      Homologous structures develop from similar embryonic substances and thus have similar basic structural and developmental patterns, reflecting common genetic endowments and evolutionary relationships. In marked contrast, analogous structures (analogy) are superficially similar and serve similar functions but have quite different structural and developmental patterns. The arm of a man, the wing of a bird, and the pectoral fins of a whale are homologous structures in that all have similar patterns of bones, muscles, nerves, blood vessels, and similar embryonic origins; each, however, has a different function. The wings of birds and those of butterflies, in contrast, are analogous structures—i.e., both allow flight but have no developmental processes in common.

      The terms homology and analogy are also applied to the molecular structures of cellular constituents. Because the hemoglobin molecules from different vertebrate species contain remarkably similar sequences of amino acids, they may be termed homologous molecules. In contrast, hemoglobin and hemocyanin, the latter of which is present in crab blood, are termed analogous molecules because they have a similar function (oxygen transport) but differ considerably in molecular structure. Corresponding similarities occur in the structures of other proteins from different species—e.g., cytochrome c and other enzymes (biological catalysts) such as the lactic dehydrogenases in birds and mammals.

Body plan and symmetry

      The bodies of most animals and plants are organized according to one of three types of symmetry: spherical, radial, or bilateral. A spherically symmetrical body is similar throughout and can be cut in any plane through the centre to yield two equal halves. A few of the simplest plants and animals are spherically symmetrical—e.g., protozoans such as Radiolaria and Heliozoia. Radially symmetrical bodies, such as those of starfishes and mushrooms, have a distinguishable top and bottom and usually have a cylindrical shape, with the body parts radiating from the central axis. A starfish can be cut into two equal halves by any plane that includes the line, or axis, running through its centre from top to bottom. The anterior, or oral, end usually contains the mouth; a posterior, or aboral, end may have an anus. In the bilaterally symmetrical body of higher animals including man, only a cut from head to foot exactly in the centre divides the body into equivalent halves. An anterior, or head, end and a posterior, or tail, end can be distinguished; and the dorsal, or back, side can be distinguished from the ventral, or belly, side. But because some internal organs of man are not symmetrical (e.g., the heart), even the right and left halves of the human body are not exactly equivalent. A few organisms—amoebas, slime molds, and certain sponges—with an irregular form, or one that changes as the organism moves, have no plane of symmetry.

Morphological basis of classification (taxonomy)

      The features that distinguish closely related species of plants and animals are usually superficial differences such as colour, size, and proportion. In contrast, the major divisions, or phyla, of the plant and animal kingdoms are distinguished by characteristics that, though usually not unique to a single division or phylum, occur in unique combinations in each.

      One morphological feature useful in classifying animals and in determining their evolutionary relationships is the presence or absence of cellular differentiation—i.e., animals may be either single celled or composed of many kinds of cells specialized to perform particular functions. Some multicellular animals have only two embryonic cell, or germ, layers: an ectoderm (outer layer) and an endoderm (inner layer), which lines the digestive tract. Other animals have these, in addition to a mesoderm, which lies between the ectoderm and endoderm. Animals may have one of two types of body cavity. The bodies of the Coelenterata (invertebrates such as the jellyfish) and other primitive many-celled animals consist of a double-walled sac surrounding a single cavity with a mouth. Higher animals have two cavities, and their bodies are constructed on a so-called tube-within-a-tube plan. An inner tube, or digestive tract, is lined with endoderm and opens at each end to form the mouth and the anus. An outer tube, or body wall, is covered with ectoderm. Between the two tubes a second cavity, or coelom, lies within the mesoderm and is lined by it. Another major distinguishing morphological feature of animal phyla is the presence or absence of segmentation. The members of several phyla have bodies characterized by the presence of a row of segments, or body units, of the same fundamental structure. Segmented animals include the vertebrates, the annelids (invertebrates such as the earthworm), and the arthropods (invertebrates such as insects); in some segmented animals such as man and most vertebrates, however, the segmental character of the body is obscured. An evolutionary tendency in many animal phyla has been the progressive differentiation of the anterior end to form a head with conspicuous sense organs and an accumulation of nervous tissues, a brain; the tendency is termed cephalization. Some morphological structures are found only in one phylum; for example, only the Coelenterata have stinging cells (nematocysts); the Echinodermata (invertebrates such as starfishes) have a peculiar water vascular system, and only the Chordates (e.g., reptiles, birds) have a dorsally located, hollow nerve cord.

      Like animals, plants may be either single celled or composed of many kinds of specialized cells. The bodies of most of the lower plants, such as algae and fungi, are comprised of the least differentiated and least specialized type of plant cells, parenchyma cells. The embryonic tissues of higher plants, unlike those of animals, remain extremely active throughout the life of the plant. In addition, the different types of cells characteristic of the body of higher plants arise from meristems, specific regions in the plant body where cells divide and enlarge. In all but the simplest forms, the plant body is composed of various types of cells associated in more or less definite ways to form systems of units called tissue systems—e.g., the vascular system consisting of conductive tissues. The arrangement of the components of the vascular system is a distinguishing morphological feature of various plant groups. The character and relative extent of the two phases in the life history of a plant—the sexual phase, or gametophyte, and the sporophyte—vary considerably among the plant groups and are useful in distinguishing them.

Areas of study

anatomy

      The best known aspect of morphology, usually called anatomy, is the study of gross structure, or form, of organs and organisms. It should not be inferred however, that even the human body, which has been extensively studied, has been so completely explored that nothing remains to be discovered. It was found only in 1965, for example, that the nerve to the pineal gland, which lies on the upper surface of the brain of mammals, is a branch from the sympathetic nerves; the sympathetic nerves receive nerve impulses from a small branch of the nerves that transmit impulses from the eye to the brain (optic nerves). Thus the pineal gland responds by a very indirect route to quantitative changes in the environmental lighting and secretes appropriate amounts of the substance it forms, the hormone melatonin.

      Detailed comparisons of the morphological features of different animals, termed comparative anatomy, provide strong arguments for the evolutionary relationships among different species. In the course of evolution, animals and plants tend to undergo adaptive morphological changes that enable them to survive under certain environmental conditions. As a result, animals only remotely related evolutionarily may come to resemble each other superficially because of common adaptations to similar environments, a phenomenon known as convergent evolution. Structural similarities—streamlined shape, dorsal fins, tail fins, and flipper-like forelimbs and hindlimbs, for example—have evolved in such varied animal groups as the dolphins and porpoises, both of which are mammals (mammal); the extinct ichthyosaurs, which were reptiles; and both the bony and cartilaginous fishes. In a like manner, the mole, an insectivore, and the gopher, a rodent, have both evolved shovellike forelimbs, an adaptation for digging.

      An opposite phenomenon, divergent evolution, occurs when animals originally closely related adapt to different environments and come to be superficially quite different. Although sea lions and seals, for example, are carnivores and thus closely related to bears, cats, and dogs, their adaptations (adaptation) to an aquatic existence have resulted in morphological characteristics distinct from those of the terrestrial carnivores. In the course of mammalian evolution, many features have changed to permit specific animal groups to adapt to particular environments—e.g., the number and shape of the teeth, the length and number of bones in the limbs, the number and attachment sites of muscles, the thickness and colour of the hair or fur, and the length and shape of the tail.

      Careful study of adaptive morphological aspects has permitted inferences about the course of the evolutionary history of various animals and of their successive adaptations to changing environments. The present-day Australian tree-climbing kangaroos (kangaroo), for example, are the descendents of a ground-dwelling marsupial, from whom evolved forms that began to live in trees and eventually developed limbs adapted to tree climbing. But the events may have occurred in the reverse sequence; that is, specialized limbs may have evolved before the animal adopted an arboreal mode of life. In any event, some of the tree-dwelling kangaroos subsequently left the trees, became readapted to life on the ground (i.e., their hindlegs became adapted for leaping), and then went back to the trees but with legs so highly specialized for leaping as to be useless in grasping a tree trunk; consequently, present-day tree kangaroos climb by bracing their feet against a tree trunk, as do bears. Careful comparisons of the feet of the many kinds of living Australian marsupials reveal the stages in this complicated process of adaptation and re-adaptation.

      Changes in genes (mutations) constantly occur and may cause a decrease in size and function of an organ; on the other hand, a change in the environment or in the mode of life of a species may make an organ unnecessary for survival. As a result, many plants and animals contain organs or parts of organs that are useless, degenerate, undersized, or lacking some essential part when compared to homologous structures in related organisms. The human body, for instance, has more than 100 such organs—e.g., the appendix, the fused tail vertebrae (coccyx), the wisdom teeth, the muscles that wiggle the ears, and the hair on the body.

      The parts of a seed plant include roots, stems, leaves, and reproductive organs in the flowers. The evolution of specialized conducting tissues called xylem and phloem has enabled seed plants to survive on land and to attain large sizes. Roots anchor the plant; enable it to maintain an upright position; and absorb water, minerals, and other nutrients from the soil. The roots of plants such as carrots, beets, and yams serve as sites for food storage. The stem links the roots with the leaves, where photosynthesis occurs, and its xylem and phloem are continuous with those of root and leaf. The stem supports leaves, flowers, and fruits. Each year, the stems of woody plants add a layer of xylem and phloem, the annual ring, the width of which varies with climatic conditions. A leaf consists of a petiole (stalk), by which it is attached to the stem, and a blade, typically broad and flat, that contains bundles, or veins, of xylem and phloem on the undersurface. The flower contains pollen-producing anthers and egg-producing ovules. After fertilization the base of the flower, or ovary, enlarges and forms the fruit, which is a mature ovary containing seeds, or mature ovules. The bodies of ferns and mosses also are composed of roots, stems, and leaves, but those of lower plants such as mushrooms and kelps are much more simple and lack true roots, stems, and leaves.

histology

      A major trend in the evolution of both plants and animals has resulted in the specialization of cells and a division of labour among them. The cells comprising a tree or a man are quite different; each is specialized to carry out certain functions. Although specialization may permit a cell to function efficiently, it also increases the interdependence of body parts; an injury to or the destruction of one part, therefore, may result in death of the whole organism. The study of the structure and arrangement of tissues (tissue), defined as groups or layers of cells that together perform certain special functions, is termed histology. Each kind of tissue is composed of cells with characteristic features such as size, shape, and relationship to adjacent cells and may also contain noncellular material—connective tissue fibres or a bony material.

      Morphologists usually separate animal tissues into six groups; epithelial (epithelium), connective (connective tissue), muscular, blood, nervous, and reproductive tissues. The cells composing epithelial tissues form a continuous layer or sheet that either covers the surface of the body or lines some cavity within the body, thus protecting the underlying cells from mechanical and chemical injury or from invasion by microorganisms. Epithelial tissues absorb nutrients and water, secrete a wide variety of substances, and may play a role in the reception of sensory stimuli. The connective tissues—bone, cartilage, ligaments, and fibrous connective tissue—support and hold together the other cells of the body. The cells of the connective tissues secrete large quantities of nonliving material (matrix), the characteristics of which largely determine the nature and the function of the specific types of connective tissue; the matrix secreted by fibrous connective tissue cells, for example, is a thick matted network of microscopic fibres surrounding the connective tissue cells. Connective tissue holds skin to muscle, keeps glands in position, makes up the tough outer walls of the blood vessels, and forms a sheath around nerve fibres and muscle cells. Tendons are flexible, cable-like cords of specialized fibrous connective tissue that join muscles (muscle) to each other or muscle to bone. Ligaments are somewhat elastic cords of specialized fibrous connective tissue that join one bone to another.

      Muscular tissues are composed of elongated, cylindrical, or spindle-shaped cells, each of which contains many small fibres called myofibrils. Muscle cells perform mechanical work by contracting—that is, by becoming shorter and thicker. The three types of vertebrate muscles include the cardiac muscle, which is found only in the walls of the heart; smooth muscles, which are found in the walls of the digestive tract and in other internal organs; and skeletal muscles, which make up the bulk of the muscle masses attached to the bones of the body. Skeletal and cardiac muscles have alternating light and dark stripes the relative sizes of which change during the contraction process. Evidence from electron microscopy indicates that two types of filaments occur in muscle; during contraction, one type of filament slides past the other.

      Nerve tissue is made of cells, called neurons (neuron), which are specialized to conduct nerve impulses. Two or more thin hairlike fibres, called axons and dendrites, extend from the enlarged cell body containing the nucleus. The neurons extending from the spinal cord to the end of an appendage (e.g., arm, leg) may extend to a metre (about three feet) or more in man and to several metres in an elephant or a whale.

      Egg cells in the female and sperm cells in the male are reproductive tissues adapted for the production of offspring. The egg cell is modified by the accumulation of considerable amounts of yolk and other food reserves. The highly specialized spermatozoon contains a tail, the beating of which propels it to the egg.

      Blood is composed of red cells, which are specialized for the transport of oxygen and carbon dioxide, and white cells, which engulf bacteria and produce antibodies (proteins formed in response to foreign substances called antigens). Blood also contains platelets, small fragments of cells from the bone marrow that play a key role in initiating the clotting of blood.

      The cells of higher plants may be differentiated into meristematic, protective, fundamental, and conductive tissues. Meristematic tissues, which are composed of small, thin-walled cells with few or no vacuoles (cavities), differentiate into the other types of plant tissue and are found in the rapidly growing parts of the plant—e.g., at the tips of roots and stems. Protective tissues are composed of thick-walled cells that protect the underlying thin-walled cells from mechanical abrasion and dehydration; examples of protective tissues include the epidermis of leaves and the cork layers of stems and roots. The fundamental tissues comprising the body of a plant include the soft parts of the leaf, the components of the pith and the cortex of stems, the roots, and the soft parts of flowers and fruits. These tissues function in the production and storage of food. Two types of conductive tissues occur in higher plants: xylem conducts water and dissolved salts, and phloem conducts dissolved organic materials such as sugars. Both types are composed of elongated cells that fuse end to end with other cells to form the sieve tubes through which substances are transported in phloem and xylem vessels.

cytology

      The living material of most organisms is organized into discrete units termed cells (cell); the study of their features is known as cytology. The cellular contents, when viewed through a microscope at low magnification, usually appear to consist of granules or fibrils of dense material, droplets of fatty substances, and fluid-filled vacuoles suspended in a clear, continuous, semifluid substance called cytoplasm. The remarkable structural complexity of the cell is more fully revealed at the higher magnifications attainable with the electron microscope; structural details of various cellular components, or organelles, as revealed by the technique known as X-ray diffraction analysis, have provided information concerning the relationships between the structures of the cellular components and of the molecules comprising them. Although most cells have certain features in common, the kinds and amounts of components vary considerably. Cellular components include structures such as mitochondria, chloroplasts, endoplasmic reticulum, Golgi complex, lysosomes, oil droplets, granules, and fibrils. The membrane around the cell is a three-layered structure called a unit membrane; similar membranes surround many cellular components—e.g., the mitochondria.

      A small spherical or oval organelle, the nucleus, is typically found near the centre of a cell. The genes within the nucleus control the development of the various traits of the cell by controlling the synthesis of specific proteins. The nuclear components are separated from those of the cytoplasm by the nuclear membrane. The structure of the nucleolus, a spherical body within the nucleus, is extremely variable in most cells. Although more than one nucleolus may occur in a nucleus, each cell of an animal or plant species has a fixed number of nucleoli. The nucleoli apparently play a role in the synthesis of the ribonucleic acid (RNA) constituent of the cellular components called ribosomes, which function in protein synthesis. Adjacent to the nucleus in the cells of animals and certain lower plants are two small, cylindrical bodies, the centrioles, which, during cell division, separate, migrate to opposite sides of the cell, and organize a structure called a spindle between them.

      Within the cytoplasm of both plant and animal cells are components called mitochondria, which may be shaped like spheres, rods, or threads. Each mitochondrion is bounded by a double membrane, the outer layer of which forms the smooth outer boundary of the mitochondrion; the inner layer, folded repeatedly into shelflike folds called cristae, contains enzymes that play an essential role in the conversion of the energy of foodstuffs into the energy used for cellular activities. The cells of most plants contain plastids, small bodies involved in the synthesis and storage of foodstuffs. The most important plastids, the chloroplasts, function in trapping the energy of sunlight during photosynthesis. They are disk-shaped structures with a platelike arrangement of tightly stacked membranes.

      The cytoplasmic components important in protein synthesis, the ribosomes, are composed of nucleic acid and protein. Clusters of five or more ribosomes, termed polysomes, appear to be the functional unit in protein synthesis.

      Lysosomes are membrane-bound structures containing a variety of enzymes that can break down the large molecular constituents of the cell. The membrane surrounding lysosomes presumably prevents the enzymes from digesting the cell contents before the cell dies.

Embryology

      The structures and the relationships among the various parts of a mature plant or animal are usually better understood if the successive developmental stages are studied. Thus, morphologists have traditionally been interested in the study of embryos and their developmental patterns—i.e., the science of embryology.

      Development typically begins in animals (embryo) with the cleavage, or division, of the fertilized egg (zygote) to form a hollow ball of cells called the blastula; the blastula then develops into a hollow cuplike body of two layers of cells, the gastrula, from which the embryo ultimately is formed. At one time, the techniques available to embryologists enabled them to study only whole embryos at different developmental stages. The science of experimental embryology began during the first half of the 20th century, when microsurgical techniques became available either for the removal and study of certain structures from tiny embryos or for their transplantation to other regions of the embryo. Advances in understanding the mechanism by which biological information is transferred in DNA and the means by which this information results in the production of specific proteins have led to efforts to describe development in biochemical terms. Although hypotheses regarding the reasons for the appearance of a specific enzyme or some other protein at a specific time during development have been formulated and tested, the biochemical basis of morphogenesis itself—that is, the reason for the development of particular structures—has not yet been established.

      The development of the seed plant is basically different from that of an animal. The egg cell of a seed plant is retained within the enlarged lower part, or ovary, of the seed-bearing organ (pistil) of a flower; two sperm nuclei pass through a structure called a pollen tube to reach the egg. One sperm nucleus unites with the egg nucleus to form the zygote from which the new plant will develop; the second sperm nucleus unites with two nuclei, called polar nuclei, to form a body called a triploid endosperm, the cells of which divide to form a nutritive mass within the seed. The zygote undergoes several cell divisions to form the embryo, which is surrounded by the endosperm. The embryo develops one or two seed leaves, or cotyledons, which may become thick and fleshy with stored foodstuffs. The epicotyl, which consists of a growing point enclosed by a pair of folded miniature leaves, develops above the point of attachment of the seed leaves. Below the seed leaves extends the hypocotyl, the tip, or radicle, of which forms the primary root of the embryonic plant. The factors involved in initiating and controlling morphogenesis in plants have been studied by growing cells, tissues, and organs derived from plants. Indeed, an entire carrot plant has been grown from one cell of a mature carrot; this provides striking evidence that the cell from the adult plant contains all of the genetic information needed to produce an entire plant, including roots, stems, and leaves. The technique of growing plants from isolated plant parts has been useful in studies involving the characteristics of embryonic growth, the correlated growth of plant parts, and the nature of differentiation and regeneration (the replacement of lost parts).

Methods in morphology

Chemical techniques

      The methods of investigating gross structure depend on careful dissection, or cutting apart, of an organism and on accurate descriptions of the parts. The study of the structure of tissues and cells has been extended by the techniques of autoradiography and histochemistry. In the former, a tissue is supplied with a radioactive substance and allowed to utilize it for an appropriate period of time, after which the tissue is prepared and placed in contact with a special photographic emulsion. Silver grains in the emulsion in contact with radioactive substances darken; thus, the location of the dark spots indicates the position at which the radioactive substance was concentrated in the tissue. Histochemistry involves the differential staining of cells (i.e., using dyes that stain specific structural and molecular components) to reflect the chemical differences of the constituents. By choosing appropriate dyes, the histochemist is able, for example, to determine the acidity or alkalinity of the chemical compounds comprising cell components. In addition, dyes that stain specific molecular constituents such as glycogen, DNA, RNA, and protein also are used. The histochemist is able to locate a specific enzyme in a thin slice of tissue, to provide the specific substance with which the enzyme reacts to form a product, and to add a compound that reacts with the product to form an insoluble coloured compound the location of which is relatively easy to determine. In this way, information has been obtained about the specific location of enzymes within the cell.

Microscopic techniques

      Histologists and cytologists utilize microscopic techniques—light microscopy (microscope), phase contrast microscopy, interference microscopy, polarization microscopy, fluorescent microscopy, and electron microscopy—to investigate certain aspects of cell structure. Phase contrast microscopy is widely used to study the structure of living cells because, with such apparatus, internal structures can be observed without killing and staining the cell. In addition, motion pictures of dividing cells or moving cells can be made using phase contrast microscopy.

      The interference microscope involves passing two separate beams of light through the specimen. With the appropriate instrument, the mass of material per unit area of the specimen can be determined, and contour mapping of small objects is possible.

      Crystalline or fibrous elements, both of which are characterized by an orderly or layered molecular structure, are studied with a polarizing microscope; the polarizing microscope has been particularly useful in studying the detailed structure of bone.

      In fluorescence microscopy, the images seen are molecules of fluorescent dyes added to cells that attach to specific cellular components. Appropriate filters are required to insure that only the light of longer wavelength contributes to the image. Fluorescent antibodies have been used to locate specific kinds of proteins and other materials in certain cells of a tissue or in certain regions of a cell. The antibodies are prepared by injecting into a rabbit an antigen (e.g., the protein myosin), which stimulates white blood cells called lymphocytes to synthesize antibodies that react specifically with the antigen. After the antibodies are isolated and purified, the fluorescent dye, fluorescein, becomes attached to them by a chemical reaction. If the fluorescent antibodies are spread over a tissue, they attach specifically to the molecules that stimulated their formation (myosin). The fluorescence microscope reveals the sites containing the antigen–antibody complex as bright luminescent areas in a dark background.

      In the scanning electron microscope, a moving spot of electrons (negatively charged particles) is used to scan an object and to produce an image similar to that which appears on a television screen. In this manner, photographs with a three-dimensional appearance can be produced. With the transmission electron microscope, a beam of electrons passes through an object, such as a cell, and is focussed on the other side onto a fluorescent screen or a photographic plate. The beam of electrons in the scanning electron microscope is focussed and then scanned across the specimen. The electrons that leave the specimen, which are not necessarily the same electrons that strike it, are then used to control the beam of a cathode-ray picture tube. Scanning electron microscopes allow photographs to be taken not only of large molecules such as DNA but of very small objects—individual atoms of elements such as uranium or thorium.

Claude A. Villee

Additional Reading

Overviews are provided by Robert D. Barnes, Invertebrate Zoology, 6th ed. (1994), an excellent modern treatment of invertebrate morphology; Peter C. Wainwright and Stephen M. Reilly (eds.), Ecological Morphology: Integrative Organismal Biology (1994); Milton Hildebrand et al. (eds.), Functional Vertebrate Morphology (1985), technical and conceptual approaches to old problems; and Claude A. Villee, Warren F. Walker, Jr., and Robert D. Barnes, General Zoology, 6th ed. (1984), a standard college-level text on vertebrate and invertebrate morphology and its relation to function. The principles governing growth and form of organisms are the subject of the classic work by D'Arcy Wentworth Thompson, On Growth and Form, 2nd ed. (1942, reissued 1992), also available in an abridged ed. edited by John Tyler Bonner (1961, reissued 1992), still a fascinating work. Renato Dulbecco, The Design of Life (1987), also treats the fundamental principles in the design of living systems. Another classic work is J.B.S. Haldane, On Being the Right Size and Other Essays, ed. by John Maynard Smith (1985). Of related interest is Knut Schmidt-Neilsen, Scaling: Why Is Animal Size So Important? (1984), on the scaling of body size and relative dimensions.Plant and animal anatomy is addressed by Katherine Esau, Plant Anatomy, 2nd ed. (1965), a beautifully illustrated text; Alfred Sherwood Romer and Thomas S. Parsons, The Vertebrate Body, 6th ed. (1986), a standard college-level text on comparative anatomy of the vertebrates; and Warren F. Walker, Jr., and Karel F. Liem, Functional Anatomy of the Vertebrates: An Evolutionary Perspective, 2nd ed. (1987), a comprehensive textbook that connects structure, function, and evolution.E.D.P. De Robertis, Francisco A. Saez, and E.M.F. De Robertis, Jr., Cell Biology, 6th ed. (1975; originally published in Spanish, 1946), is an excellent text describing the cytological features of animal and plant cells. Don W. Fawcett, The Cell, 2nd ed. (1981), collects superb electron micrographs of several kinds of cells and cell organelles, with a brief description of each.General studies focusing specifically on embryology include Bruce M. Carlson, Patten's Foundations of Embryology, 6th ed. (1996), a mechanistic approach to chick and human development; Jan Langman, Langman's Medical Embryology, 7th ed. by T.W. Sadler (1995), on human development and congenital anomalies; Scott F. Gilbert, Developmental Biology, 4th ed. (1994), contemporary developmental biology from molecule to organ; J.M.W. Slack, From Egg to Embryo: Regional Specification in Early Development, 2nd ed. (1991), focusing on concepts, theory, and future directions; and Jonathan Bard, Morphogenesis: The Cellular and Molecular Processes of Developmental Anatomy (1990), emphasizing the many unanswered questions. The short book by Lewis Wolpert, The Triumph of the Embryo (1991), is intended for the nonspecialist reader, yet it includes sufficient information to provide an account of the major themes of the subject with modern flavour. David De Pomerai, From Gene to Animal: An Introduction to the Molecular Biology of Animal Development, 2nd ed. (1990), also conveys the main ideas on the subject as of its publication date. D.M. Glover and B.D. Hames (eds.), Genes and Embryos (1989), focuses principally on early events in development, especially the genetic control of embryogenesis, during which time the fates of most cells become established. T.J. Horder, J.A. Witkowski, and C.C. Wylie (eds.), A History of Embryology (1986), provides a chronicle of history, data, and concepts from the early 19th century through the origins of molecular genetics.Works on embryology at a more advanced level are Leon W. Browder (ed.), Developmental Biology: A Comprehensive Synthesis, 7 vol. (1985–91), treating a diversity of topics from molecule to organ; Brian K. Hall, Evolutionary Developmental Biology (1992), a history and analysis of concepts; and Michael Akam et al. (eds.), The Evolution of Developmental Mechanisms (1994), on the development, paleontology, and evolution of living organisms. Gerald M. Edelman, Topobiology: An Introduction to Molecular Embryology (1988), discusses general principles of spatial and temporal organization of living cells with reference to their molecular basis.Claude A. Villee Ed.

▪ linguistics

      in linguistics, study of the internal construction of words. Languages vary widely in the degree to which words can be analyzed into word elements, or morphemes (q.v.). In English there are numerous examples, such as “replacement,” which is composed of re-, “place,” and -ment, and “walked,” from the elements “walk” and -ed. Many American Indian languages have a highly complex morphology; other languages, such as Vietnamese or Chinese, have very little or none. Morphology includes the grammatical processes of inflection (q.v.) and derivation. Inflection marks categories such as person, tense, and case; e.g., “sings” contains a final -s, marker of the 3rd person singular, and the German Mannes consists of the stem Mann and the genitive singular inflection -es. Derivation is the formation of new words from existing words; e.g., “singer” from “sing” and “acceptable” from “accept.” Derived words can also be inflected: “singers” from “singer.”

* * *