Symbiosis: Rich, Exciting, Neglected Topic
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symbiosis: Rich, Exciting, Neglected Topic JANE THOMAS ROWLAND benefit by the association but relations are not obli- gatory; he reserves the term "mutualism"for asso- ciations in which both populationsare not only bene- fited but cannot survive under natural conditions without each other (obligate symbiosis). As early as 1876 the term "commensalism"was used by Van Beneden to describe the relationship between what he otherwise called "messmates." He states, "The messmate does not live at the expense of his host; IN ECOLOGIC STUDIES of biotic factors an approach all that he desires is a home or his friend's super- through the phenomenon of symbiosis and interde- fluities" (Van Beneden 1876:1). Downloaded from http://online.ucpress.edu/abt/article-pdf/36/2/77/31514/4444655.pdf by guest on 26 September 2021 pendence of organisms can be most meaningful and Because true parasitism is treated extensively in enlightening to the student at any educational level. many books and journals, it is assumed here that this It is my opinion that symbiosis has been greatly relation gets its due credit; so I will concentrate on neglected and underemphasized in general-biology symbiosis in a more narrow sense, which includes textbooks, as well as in much of the current ecologic various degrees of mutualism and commensalism, literature and the environmental studies that are so together with the relationship often called "social popular today. Many textbooks devote only a few parasitism." In social parasitism the symbiont does paragraphsto symbiosis, and these usually resort to not live in or on its host and does not receive its the most commonly used examples, leaving the im- nourishment from the tissues of its host. The social pression with the student that there are only these parasite exploits the host organism in other ways, few situations and that symbiosis is an exception such as stealing some of its food, using it as a slave, rather than a usual occurrence in animal behavior or seeing that it incubates its eggs and raises its and in plant-and-animalassociations. The purpose of young. "Behavioral symbiosis" is a term used to de- this article is to encourage the presentation of a scribe associations in which the contact may not be wider selection of examples to students and to show intimate or persisting but involves certain behavioral the universal principle of interdependence of spe- patterns of the organisms involved. cies, which takes on varying gradients of what can easily be categorized as symbiosis. Importance of the Study of Symbiosis Definitions of Terms The interdependence of organisms is a most out- standing principle of nature, whether it is at a micro- There is considerablecontroversy as to what should scopic level and involves nutritional exchange at the and should not be classed as symbiosis. The term tissue or cellular level or involves occasional contact symbiosis (Greek, "living together") was first pro- or behavior at the organismiclevel. No organism can posed by DeBary in 1879 to refer to organisms of exist in nature without others in its environment. two species living together. DeBary used lichens as These dependencies include the need for food, habi- a representative example of symbiosis; so it is quite tat, protection,reproduction, companionship, or other logical that the most popular usage of the term refers factors or a combination of these. The fascinating to mutually beneficial relationships. However, the aspect is how different organisms meet their needs broad meaning includes all intimate associations; in such diverse ways. It is the diversity through evo- therefore symbiosis can be divided into three main lutionary adaptation that presents the student with categories: (i) mutualism, in which each partner such a variety of unusual examples. However, one receives benefit; (ii) commensalism, in which one partner (the commensal) is benefited but the host The author is assistant professor of biology, is unaffected; and (Miii)parasitismn, in which the host Paine College, 1235 15th St, Augusta, Ga. is exploited to the benefit of the parasite (symbiont). 30901. She is a 1952 graduate of Tift College Needless to say, no distinct boundaries separate and has her M.S. from Emory University. Before joining the Paine faculty she taught these associations. This fact complicates defining and at Armstrong State College, Savannah, 1953- classifying the many associations of organisms.Inter- 56, and at Augusta College, 1961-84. She actions between species may be very complex, and frequently contributes book reviews to additional terms are applied to specific relationships. Choice, a publication of the Association of Authorities differ in their usages of such terms. College and Research Libraries. Library research of the kind reflected in this paper has been a primary interest of Odum (1971:211) uses the term "protocooperation" the author for the past 10 years. Mrs. Rowland's husband is to designate an association in which both populations the librarian of Augusta College. 77 does not have to search out the unique (however Other examples of cleaning activity include ground exciting this may be) to find examples of symbiosis. finches removing ticks from the skin of tortoises Every habitat, every community of plants and ani- (MacFarland and MacFarland 1972:632); a lark mals is well supplied with many symbionts. From pecking parasites from the back of the imouran, a the lichens on barren rocks or intestinal bacteria in large rodent of Mongolia (Pearse 1939:513); red the human body to the flowers and insects in the crabs of the Galapagos Islands eating ticks from garden, a student can witness the marvelous inter- the leathery skin of iguanas (Simon 1970:96; Feder action of species. 1966:329); a pseudoscorpion cleaning a lizard (Feder In this day, when molecular biology is so popular, 1966:329); mites cleaning dung deposits from the there has been a definite decrease in interest in cer- body of a dung beetle (Burton 1969:52); pseudo- tain aspects of natural history. This has occurred in scorpions removing mites from long-horned beetles spite of the soaring interest in environmental biology (Dudley 1965:49; Simon 1970: 80-83); and barber and is evidenced by a relative decline in the amount ants grooming their host ants (Simon 1970:83). of information published on symbiosis, except in Many commensal organisms live in the nests or microbiology (nitrogen fixation and the like). Be- burrows of their hosts and are cleaners or scaven- havioral symbiosis, such as is evidenced in birds gers, removing unwanted scraps of food, parasites, and in insects, was a popular topic in many works and excrement. The clownfish, in addition to its pro- published about 1900-1950. Some of the most thor- tective association with anemones (mentioned be- Downloaded from http://online.ucpress.edu/abt/article-pdf/36/2/77/31514/4444655.pdf by guest on 26 September 2021 ough works are now quite dated, and there are low), has been observed grooming the tentacles of relatively few books in print that deal exclusively the anemone. with symbiosis. The whole organism and its relation to its environment should still receive attention. Beater Associations Fortunately, there are periodicals that still deal with A beater, or driver, is an animal that disturbs the natural history of organisms, including the biotic small creatures as it moves about; and other animals, factors of environment; among these are Natural such as insectivorous birds, prey upon the animals History, Audubon Magazine, and Animals. flushed. Rand (1954) cites descriptions of a number of such relationships. A common example is the asso- Symbiosis Takes Many Forms ciation of cattle egrets with large herbivorous mam- mals (Peterson 1954). The carmine bee-eater uses Because there are so many types of symbiosis and the kori bustard as its beater; and, as is the case such a large number of close interdependencies of with many of these symbionts, it often rides on the different species, this paper will emphasize only five back of the beater, frequently jumping off to capture categories, which can serve to show the wide hetero- insects (Rand 1954). Pompadoured hornbills, or specific situations represented. These categories are monkeybirds, in Africa associate with guenon mon- cleaning symbiosis, beater associations, pollination keys, feeding on the insects made more easily avail- symbiosis, symbiosis for shelter or protection, and able to them (Burton 1969:36; Simon 1970:31). One social parasitism. species of African hornbill follows driver-ant col- umns, before which small animals are fleeing (Simon Cleaning Symbiosis 1970:32). Other birds also associate with driver ants In the study of symbiosis many aspects of animal for this purpose; for example, the antbirds (Willis behavior are observed. Cleaning symbiosis, for ex- 1973). Welty (1962:380) says the rough-legged hawk ample, involves a wide variety of organisms. "Clean- follows the Arctic fox, to feed on the mice it stirs up. ing symbiosis is defined as a relationship during In the U.S. we often see cattle accompanied by cow- which certain organisms, known as cleaners, remove birds, which gain from the association by the greater ectoparasites, bacteria, diseased and injured tissues, number of insects they are able to catch. and unwanted food particles from cooperating hosts" (Cheng 1970:26). This has been studied most thor- Pollination Symbiosis oughly in marine organisms (Limbaugh 1961; Feder Interdependence of plants and animals becomes 1966). Certain cleaner organisms, such as fish and most interesting and varied with pollination ecology. shrimp, set up cleaning stations and have an eager Whenever a plant is dependent upon a biotic agent clientele of fishes waiting to be cleaned of parasites, for its pollination, this approximates symbiosis. Some diseased tissue, and debris. The host is benefited, plants, however, have developed such great specific- and the cleaner is rewarded with a meal of the clean- ity in the methods of pollen transfer that they and ings.