SEEKING to UNDERSTAND the LIVING BIRD the 1997 Margaret Morse Nice Lecture

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SEEKING to UNDERSTAND the LIVING BIRD the 1997 Margaret Morse Nice Lecture Wilson Bulletin, 110(l), 1998, pp. 28-44 SEEKING TO UNDERSTAND THE LIVING BIRD The 1997 Margaret Morse Nice Lecture NICHOLAS E. COLLIAS.2’ AND ELSIE C. COLLIAS ’ For over 50 years the classic study by Mar- bird, an impression still fresh after three-quar- garet Morse Nice on the life and behavior of ters of a century. I was not then aware that the Song Sparrow (Melospiza melodia) has anyone in my world had any real interest in given a basic orientation and ideas for re- birds aside from myself and Peter Rabbit, the search to investigators of the natural behavior hero of the bedtime stories by Thornton W. of birds. She made the first detailed, long-term Burgess. It was far from my thoughts that any- study of the behavior of wild birds color- one could make a living watching birds, until banded for individual identification (Nice years later when a scholarship to the Univer- 1933, 1934). She had an excellent education sity of Chicago made possible for me a career and the two volumes of her monograph on the in zoology and ornithology. Song Sparrow (Nice 1937, 1943), one on pop- The situation was different for Elsie, some ulations, one on behavior, are written from the of whose ancestors, like those of Margaret viewpoint of basic biological problems, enliv- Nice, arrived in New England in the 1600s. ened by her profound curiosity about the life Eventually, Elsies’ family settled in Tiffin, of birds. Mrs. Nice was a firm believer in con- Ohio, near the Sandusky River. Her parents firming for herself any important published encouraged her early interests in nature and fact about bird behavior, which is the reason taught her the local birds. Her father, Heath why research was a passion with her (Nice K. Cole, was a lawyer with a strong interest 1979). In this spirit of direct scientific inquiry, in the natural sciences. Elsie, before she start- we wish to present some highlights of our ed school, developed a great interest in insect own research. life, and readily prevailed on her father to We have spent much of our lives trying to make her an insect net. She eventually donat- understand the biology of avian behavior, es- ed a substantial insect collection to Heidelberg pecially the function, causes, ontogeny and College, her alma mater in Tiffin. After a year evolution of social behavior at the physiolog- in research on malarial mosquitoes with the ical, individual and population levels of or- United States Public Health Service, Elsie ganization (N. Collias 1991). The first part of went back to finish her Ph.D. work in zoology this presentation will be given by Nicholas, at the University of Wisconsin, where we met. the second by Elsie, but we have generally Our friendship included getting up well before worked together. dawn to watch the prairie-chickens dance near Let me briefly introduce ourselves. My ear- Plainfield, Wisconsin, where we were two of ly childhood was in Chicago Heights, then a the thousands of guests that Fran and Fred small town, where my family lived on the Hamerstrom introduced to these fascinating edge of town. Soon after learning to read, I birds over the years. After Elsie and I were discovered in the local library “The Burgess married we did our research together on birds. Bird Book for Children” (Burgess 1919) with UNIVERSITY OF CHICAGO (1937-1942). its magnificent color plates by Louis Agassiz HORMONES AND BEHAVIOR Fuertes. These plates gave me names for the common birds that I encountered in the nearby Mrs. Nice and her family moved to Chicago fields and woods. I vividly recall my first in 1936 and bought a house near the Univer- glimpse of a bluebird in an orchard. It seemed sity of Chicago where I was a student, and we as if a fragment of the blue sky had fallen and became friends. was flying about on the back and wings of a My research career began after graduation in 1937 when Professor W. C. Allee of the ’ ’ Univ. of California, Dept. of Biology, Los Angeles, University of Chicago zoology department CA 900951606. hired me as his research assistant to work on * Corresponding author. the effects of hormones on the peck order of 28 Collias and CoZZiasl THE LIVING BIRD 29 SYNCHRONY - GONADOTROPHIN - SEXUAL BEHAVIOR general way with more current evidence as OF INTERNAL 4 f CYCLES WITH critically reviewed in 1996 by Buntin, who ENVIRONMENT I-) I /G0NAGSi also brings out the variation between different PROLACTIN _ PARENTAL BEHAVIOR species of birds. FIG. I. Interactions between hormones and behav- There are two general phases in the breed- ior in the control of breeding behavior in doves and ing cycle of birds, a sexual phase stimulated pigeons. Inhibition indicated by minus (-) sign. (After by pituitary gonadatrophin and gonadal ste- N. E. Collias 1952 and Collias and Collias 1984). roid hormones, and a parental phase dominat- ed by pituitary prolactin. The gonadal hor- mones stimulate pair formation, nest building hens. Testosterone and estradiol had been iso- and copulation. Male Ring Doves stimulate lated in pure form in 1935 (Martian 1950). In the females to lay eggs, as I found in a con- chickens, the cocks’ testes secrete testoster- trolled experiment (N. Collias 1950), confirm- one, the hens’ ovary secretes testosterone, es- ing Wallace Craigs’ demonstration of the same tradiol, and progesterone (reviewed in Collias thing in 1911. Gonadal hormones, together and Collias 1984). with the mate, nest and eggs, also help bring We found that ovariectomized hens injected about onset of incubation. with testosterone became more aggressive and In 1935, Oscar Riddle and his colleagues rose in the peck order, some from the bottom discovered that prolactin injected into laying to the top (Allee et al. 1939). Ovariectomized hens stimulates maternal behavior, i.e., incu- hens injected with the female sex hormone, bation and care of chicks. Sitting on eggs in estradiol, became sexually receptive but they turn stimulates prolactin secretion. However, did not rise in the peck order (Allee and Col- a hypophysectomized male pigeon that I in- lias 1940). jected with estradiol thereupon paired with a Rank in the peck order is decided by fight- normal female and helped her incubate her ing or passive submission at the initial en- eggs for over a month, showing that prolactin counter of each pair of birds. Success in initial is not absolutely necessary for parental be- encounters between normal hens is correlated havior (N. Collias 1950), although it normally with larger comb size, a specific indicator of stimulates it (Buntin 1996). By inhibiting go- testosterone. In my Ph.D. thesis, using the nadotrophin, rising prolactin levels help end path coefficient method of Professor Sewall the sexual phase of breeding. Wright and with his advice, I found that a Today, comparative endocrinology of wild much greater percentage of the variance in birds in nature is a relatively new and flour- success was determined by differences in ishing field of research, made possible by such comb size than in body weight or in previous techniques as radioimmunoassay of minute success, while molt had a depressing effect on hormone levels in the blood, a technique for success (N. Collias 1943). which Rosalyn Yalow of New York received Mrs. Nice, who was president of the Chi- a Nobel Prize in 1977. For many species of cago Ornithological Society from 1939 to birds it has now been amply confirmed that a 1941, invited me to speak before that society, sexual phase dominated by steroid sexual hor- and I spoke about hormones and behavior of mones is succeeded by a parental phase more birds as a self-regulating system with positive or less dominated by prolactin (Buntin 1996), and negative feedback relations. Ten years lat- for example, in the Song Sparrow (Wingfield er (N. Collias 1950), I reviewed the literature and Goldsmith 1990). on the subject in some detail. Figure 1 shows the interactions between the UNIVERSITY OF WISCONSIN environment, hormones, and behavior. Origi- (1947-1951). ANALYSIS OF FAMILY nally it was based largely on pigeons and INTEGRATION AND VOCAL doves (N. Collias 1952), but indications now COMMUNICATION are that it may apply to many other birds as I served as Instructor in Zoology at the Uni- well. This figure was confirmed by a review versity of Wisconsin for four years. After 3 of more recent literature in 1984 (Collias and and l/2 years in the Army in World War II, I Collias 1984), and I think is consistent in a was more interested in family life than in peck 30 THE WILSON BULLETIN * Vol. IZO, No. I, March 1998 orders. Like Mrs. Nice, who as a young girl (3-4 days) some specific individuals may lead raised chickens, my first acquaintance with the other chicks to maternal clucking (from a details of individual bird behavior was with speaker) or to a source of warmth (warm chickens. The focus of my research at Wis- lamp) in a cool room. A chick that forges very consin was on family integration, and this led far ahead of the others, however, may utter to an interest in vocal communication. distress cries and is likely to turn back to re- Family integration and leadership.--When join its companions. A good leader has both a baby chick is lost, cold, hungry, thirsty, or independence, and empathy with companions in pain, it utters distress cries (chirps). When or young ones (N. Collias 1952). the chick is returned to its mother or to its Social guidance by the parent bird or other siblings, or the specific distress is relieved, the individuals enters into every stage of social chick promptly switches to pleasure notes development, and climaxes when birds learn (twitters).
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