There are many advantages to studying birds; for example, they exhibit a wide range of social behaviors, can be easily observed in both the laboratory and field settings, and have neuroendocrine systems that share fundamental properties with other vertebrate species including mammals Konishi et al. These investigators have used a variety of animal species in their investigations Adkins-Regan ; Marler ; in particular, ethologists and comparative psychologists interested in the study of the mechanistic control of behavior came from a tradition in which birds were widely used Marler The study showed a remarkable change in the behavior of the males: The basic argument of those advocating the functional approach to Pavlovian conditioning is that natural stimuli are those that would ordinarily predict a functional outcome e. Beach and colleagues began studying the behavioral endocrinology of Japanese quail in the s Beach and Inman ; Sachs , , and in the s Adkins-Regan established basic facts about the hormonal control of male reproductive behavior in quail Adkins , ; Adkins and Adler In conditions that predict the arrival of the female, males also released a greater volume of sperm and greater numbers of spermatozoa than control subjects mating with the female in a new environment Domjan et al. In the s Domjan and Hall identified a form of associative learning that they named learned social proximity response.
We developed a faster version of this test in which the proximity response is quantified during a 5-minute period of presentation of the female after the male has been habituated to the test chamber Figure 1. Quail have also been used to establish the effects of early experience on mating preference; for example, males prefer slightly unfamiliar females to females with which they were raised Bateson The entire test can be completed in 25 minutes as opposed to the procedure used by Domjan and collaborators in which birds were tested throughout the day and thus makes it possible to quantify appetitive sexual behavior in a large number of subjects each day see Balthazart et al. Although the value of this distinction has its critics Sachs , it is useful for the mechanistic analysis of male sexual behavior and other rewarding activities Ball and Balthazart More recent studies have identified more complex effects of the early social environment. Conditioned responses are relatively easy to achieve in quail through Pavlovian conditioning procedures using sexual reinforcements Domjan et al. Quail have also been used in sexual conditioning studies illustrating how natural stimuli predict successful reproduction via Pavlovian processes. However, natural stimuli are not arbitrary given that individuals have experience with them already, whereas most CS in traditional Pavlovian studies are selected because they are arbitrary and therefore neutral for the animal Domjan One study showed that male quail are more responsive to minimal cues from females if these are presented in a context previously paired with access to a female Hilliard et al. If one wants to study how learning facilitates natural rewards one should not expect the most important stimuli to be neutral. The study showed a remarkable change in the behavior of the males: The fact that natural stimuli facilitate functional outcomes in quail is evident from greater reproductive success after sexual learning. Male sexual behavior has arousal and satiety components, comparable to the appetitive and consummatory aspects of behavior articulated by ethologists such as Niko Tinbergen, Konrad Lorenz, and Gerard Baerends and even earlier based on the pioneering work of Wallace Craig, among others see Balthazart and Ball b. The origins of behavioral endocrinology are truly interdisciplinary in that the field attracted investigators trained in the behavioral sciences such as experimental psychology or ethology as well as those trained in endocrinology or neurobiology Beach ; Marler This effect was observed only if both males and females had been conditioned but not if only one member of the pair associated the CS with sexual interactions Mahometa and Domjan His group showed that long photoperiods are necessary to induce full gonadal development in quail and that photoperiodic experience can render the birds nonresponsive to photoperiods that previously stimulated reproductive development Robinson and Follett This change in preference is not dependent on the fact that the male has had additional copulatory experience that might in turn change his behavior in such a way as to make him more attractive to the female. This differential brain activation could represent at least a component of the mechanism that mediates improved copulatory efficiency associated with the presentation of a CS before sexual interactions. The enzyme aromatase converts testosterone to estradiol and is enriched in the POM in a male-biased fashion. In the s Lehrman , took the lead in illustrating the value of birds to studies of the hormonal control of reproductive behavior, based on his now classic studies on ring doves, and Hinde conducted parallel studies on canaries. These investigations formed one of the threads that has led to the field endocrinology movement spearheaded by Wingfield and colleagues, who have investigated hormone-behavior relationships in a wide range of avian species in their natural environments Wingfield ; Wingfield and Silverin See other articles in PMC that cite the published article. These conditioned stimuli CS 1 were then used, when presented alone, to modify aspects of male sexual behavior; for example, in one study such a CS elicited the strutting display that males normally exhibit toward females Farris There are many advantages to studying birds; for example, they exhibit a wide range of social behaviors, can be easily observed in both the laboratory and field settings, and have neuroendocrine systems that share fundamental properties with other vertebrate species including mammals Konishi et al. Appetitive behaviors are an expression of sexual motivation, and the functional outcome of stereotyped consummatory behavior is a reduction in motivation Timberlake and Silva They can be easily bred in captivity with the help of an incubator.
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