morphology
morphology, in biology, the study of the size, shape, and structure of animals, plants, and microorganisms and of the relationships of their constituent parts. 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 comprised by the bodies of animals and the roots, stems, leaves, and flower parts comprised by 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 of 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 humans appreciated the form and structure of their 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 Middle East, as humans learned to domesticate certain animals and to cultivate many fruits and grains, they 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 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, 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, 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 English physician William Harvey, 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, Dutch microscopist Antonie van Leeuwenhoek 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, who was associated with morphology in all its aspects, remains a classic in the field. British surgeon John Hunter and French zoologist Georges Cuvier 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, 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 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 focused 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 are superficially similar and serve similar functions but have quite different structural and developmental patterns. The arm of a human, 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, and 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 described as 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 Heliozoa. 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 humans, 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 humans 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
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 humans 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 called 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, comprise 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.
