Progress 10/01/11 to 09/30/16

Outputs
Target Audience:Applied ethologists, animal behaviorists, animal scientists and livestock producers. Changes/Problems: Nothing Reported What opportunities for training and professional development has the project provided?FourPhD students completed training on this project. Five undergraduate students completed senior honors theses and scores of undergraduate assistants learned animal handling techniques, research methodologies, and behavioral hypothesis testing as a result of this project. How have the results been disseminated to communities of interest?Poster presentations on phases of this project were presented to the scientific community each year of the project. What do you plan to do during the next reporting period to accomplish the goals? Nothing Reported

Impacts
What was accomplished under these goals? Due to the differential cost of reproduction in promiscuous species, like domesticated goat(s) (Capra hircus), it is expected that females should mate with higher quality males, while males should mate with a greater number of females. Females may use conspicuous secondary sexual characteristics of males such as courtship display to distinguish among high and low quality males. Our studies suggest that courtship rate is T-dependent. Further, females can use courtship rate to distinguish among males. Preference testing has shown that sexually experienced male goats choose females that are tail wagging, a behavior that may function as both attractivity and proceptivity, over those that are not. We hypothesized that exposure to females expressing high rates of tail wagging would arouse males, increasing sexual performance. Our studies supported the hypothesis that male goats are sexually aroused by tail wagging. This study and previous work demonstrate that tail wagging functions as both attractivity and proceptivity in goats. In a promiscuous species like the domestic goat (Capra hircus), in which maternal investment is greater than paternal investment, a female may mate selectively with a more-fit male to improve her reproductive fitness. Several studies were conducted to examine if females use testosterone-dependent cues to assess potential mates. We demonstrated that females preferred the males with higher testosterone concentrations and we concluded that females use testosterone-dependent cues to assess potential mates. Moreover, circulating testosterone concentrations may indicate a male's overall fitness. We examined testosterone concentrations in male goats exposed to different reproductive opportunities. It was hypothesized that reproductive readiness would be stimulated by proximity to females. Thus, we predicted that testosterone concentrations would be higher for males housed in the presence of females. We also examined the relationship between testosterone concentrations and body mass for males housed in the presence or absence of females and for testosterone-treated castrated males during the breeding season. Further, we hypothesized that body condition would be directly influenced by circulating testosterone concentrations and thus, it was predicted that males with higher T concentrations would experience increased weight loss during the breeding season. In summary the results of these studies demonstrated that maintenance of high testosterone concentrations is costly (body mass loss). Importantly, variation in testosterone concentrations provides males with the ability to maximize reproductive success by modifying energetic costs depending on reproductive opportunities available.

Publications

• Type: Journal Articles Status: Published Year Published: 2016 Citation: Longpre K.M., Guterl, J.N., and L.S. Katz. 2016. Proximity to females alters circulating testosterone concentrations and body weight in male goats. Small Ruminant Research 144:334-340. 10.1016/j.smallrumres.2016.10.014
• Type: Journal Articles Status: Published Year Published: 2014 Citation: Longpre, K.M., Kinstlinger, N.S., Mead, E.A., Wang, Y., Thekkumthala, A.P., Carreno, K.A., Hot, A., Keefer, M., Tully, L., Katz, L.S., and Pietrzykowski, A.Z. 2014. Seasonal variation in urinary microRNA expression in male goats (Capra hircus) as assessed by next generation sequencing. General and Comparative Endocrinology 199:1-15. 10.1016/j.ygcen.2014.01.002
• Type: Conference Papers and Presentations Status: Published Year Published: 2016 Citation: Fritz, W.F., Becker, S.E. Alexander, B.M. and L.S. Katz. 2016. Simulated Self-enurination Increases Serum Luteinizing Hormone Concentrations in Male Goats (Capra hircus). Society for the Study of Reproduction, San Diego, CA.

Progress 10/01/14 to 09/30/15

Outputs
Target Audience:Animal scientists and behavioral endocrinologists. Changes/Problems: Nothing Reported What opportunities for training and professional development has the project provided?Undergraduate animal science students learned how to conduct behavior studies, collect and analyze data and present findings of their studies in written and oral formats. How have the results been disseminated to communities of interest?Yes, presentation was made to members of the Society for Behavioral Endocrinology (Fritz, W.F., Sena, L.S, Becker, S.E. and L.S. Katz. 2015. Differential effects of androgens and estrogens on sexual behaviors in the male goat. (Capra hircus). Society for Behavioral Neuroendocrinology, Asilomar, California.) A presentation was also made to the Rutgers University Endocrine Programs's annual Pioneers in Endocrinology workshop. What do you plan to do during the next reporting period to accomplish the goals?We will continue to examine the impact of self-enurination on the reproductive endocrinology of male goats.

Impacts
What was accomplished under these goals? The endocrinology of self-enurination (SE), a sexual behavior exhibited by male goats, is poorly understood. Twenty castrated male goats (wethers) were randomly divided into five groups and injected for seven weeks with one of the following: 25 mg testosterone propionate (TP), 25 mg dihydrotestosterone propionate (DHTP), 100 μg estradiol benzoate (EB), 100 μg EB and 25 mg DHTP (EB+DHTP), or oil (CON). The effects on SE were assessed by using two 10-min behavior tests we have shown to promote SE. In one test, a wether watched (from 4.6 m) a buck courting and mounting an estrous female. In the other, he watched a buck that could only court the estrous female through a barrier. Finally, to record the effect of treatment on courtship and mounting in addition to SE, each wether was placed loose in a pen with an estrous female for 10 min. After a five-week, treatment-washout period, wethers were randomly assigned to different treatment groups and tested again. Courtship frequencies displayed in all three scenarios were greater than CON only for groups exposed to estrogen (TP, EB+DHTP, EB). Self-enurination frequencies exhibited during behavior tests in which the wether was watching were greater than CON only for androgen-treated groups (TP, EB+DHTP, DHTP). In contrast, when the wether was free to interact with the female only the DHTP group displayed SE at a higher frequency than CON. Treatment had no effect on mount frequencies in this test scenario, however ejaculation frequencies were highest for TP and EB+DHTP. These studies suggest that the social context of the behavior tests contributes as much to behavior expression as steroid treatment.

Publications

• Type: Conference Papers and Presentations Status: Published Year Published: 2015 Citation: Fritz, W.F., Sena, L.S, Becker, S.E. and L.S. Katz. 2015. Differential effects of androgens and estrogens on sexual behaviors in the male goat. (Capra hircus). Society for Behavioral Neuroendocrinology, Asilomar, California.

Progress 10/01/13 to 09/30/14

Outputs
Target Audience: Nothing Reported Changes/Problems: Nothing Reported What opportunities for training and professional development has the project provided? One PhD student led this research effort. In addition many undergraduate Animal Science students were given the opportunity to participate in the research, helping with animal handling and behavioral data collection. How have the results been disseminated to communities of interest? A poster presentation was delivered to scientific members of the Animal Behavior Society and participants in the Rutgers University Annual Endocrine Workshop. What do you plan to do during the next reporting period to accomplish the goals? Behavioral and endocrine affects of chemical signals carried in goat urine will be measured.

Impacts
What was accomplished under these goals? Self-enurination (SE), self-marking with urine, in male goats occurs frequently during the breeding season. This behavior may present attractive olfactory cues to females. SE may also serve as a self-excitatory sexual behavior, in which the goat uses chemical cues to stimulate his HPG-axis via the accessory olfactory system. Our lab has shown that mature bucks had increased serum testosterone (T) concentrations in response to simulated SE, but T concentrations were variable. Measuring luteinizing hormone (LH) instead might reduce this variability. During the breeding season bucks (n=13) were allowed to court estrous females, after which they were pulled away. SE urine was then collected and frozen for use in this study, conducted in the breeding season. Each buck received 3 mL of his own SE urine or saline, one week apart in a switchback design. Blood was drawn at -10, -5, 0, 5, 10 15 and 20 min, relative to treatment. Whereas treatment with SE urine resulted in more time spent in flehmen (p<0.05) than saline (44 sec; n=13 vs. 20 sec; n=10), serum LH concentrations did not differ between treatments. Cues from SE may act as a releaser rather than a primer pheromone.

Publications

• Type: Conference Papers and Presentations Status: Published Year Published: 2014 Citation: Simulated self-enurination induces flehmen but does not affect serum LH concentrations in male goats. Animal Behavior Society, Princeton, NJ.

Progress 10/01/12 to 09/30/13

Outputs
Target Audience: Scientific community in the area of animal behavior, reproduction, and evolutionary biology. Changes/Problems: Nothing Reported What opportunities for training and professional development has the project provided? A graduate student was trained in behavioral science and endocrinology. Undergraduate students were provided training in the conduct of animal behavior studies. How have the results been disseminated to communities of interest? Presentation at annual meeting of the society for behavioral neuroendocrinology. What do you plan to do during the next reporting period to accomplish the goals? We are examining the effects of self-enurination (SE) in male goats on the neuroendocrine axis of the animal displaying this behavior. The hypothesis being tested is that SE up-regulates the animal's own reproductive axis by stimulating LH secretion.

Impacts
What was accomplished under these goals? Self-enurination (SE), self-marking with urine, in male goats occurs frequently during the breeding season and only occasionally during the nonbreeding season. This behavior may present attractive olfactory cues to females. It was hypothesized that SE may also serve as a self-excitatory sexual behavior. To reliably elicit SE, a preliminary study was conducted in which bucks were placed in various social scenarios and frequency of SE was recorded. Sexually aroused males denied access to females displayed SE. Self-enurination urine was collected from bucks (n=12; 2-9 yr of age) during the breeding season. Urine was stored frozen for use in the current study, conducted outside of the breeding season. In a switchback design, each buck was treated with a spray directed at his snout of either 1.5 ml of his SE urine or 0.9% saline on separate days. Blood was drawn at -20, 0, 20, 40, 60 and 80 min., relative to treatment, for later testosterone analysis. Treatment with SE urine was more likely than saline to elicit the flehmen response (11/12 vs. 4/12 bucks, respectively; p=0.003). Serum concentrations of testosterone were higher at 40, 60 and 80 min. (1.3, 1.6, and 1.5 ng/ml, respectively) than at -20, 0, and 20 min. (0.5, 0.5, and 0.6 ng/ml, respectively) regardless of treatment, (p<0.05). In the sexually and socially mature bucks (older than 3 yr of age; n=8), testosterone was higher in response to SE urine than to saline at 60 and 80 min. (2.2 and 2.2 vs. 1.4 and 1.1 ng/ml, respectively; p<0.05). These preliminary findings suggest that mature bucks frequently display SE during the breeding season to increase testosterone for reproductive success.

Publications

• Type: Conference Papers and Presentations Status: Published Year Published: 2013 Citation: Simulated self-enurination in male goats (Capra hircus) induces flehmen response and increases serum testosterone concentrations. Society for Behavioral Neuroendocrinology, Atlanta, GA.

Progress 10/01/11 to 09/30/12

Outputs
OUTPUTS: Studies of female mate choice in goats were continued. Findings were presented at the annual meeting of the Society of Behavioral Neuroendocrinology in Madison, Wisconsin. Manuscripts are in preparation for peer review. PARTICIPANTS: Susan E. Becker, MS., was the research assistant participating in all phases of the experiments. William F. Fritz, Jr., pre-doctoral student, conducted all of the behavior experiments. In addition, dozens of undergraduate animal science students assisted with animal handling and data collection. TARGET AUDIENCES: Scientific community in the area of Animal Behavior, Reproduction, and Evolutionary Biology. PROJECT MODIFICATIONS: Nothing significant to report during this reporting period.

Impacts
One goal of this research was to demonstrate that the domesticated female goat (Capra hircus)may be used as a model for the study of female mate choice. Implications for this research may extend to a variety of other ungulate species that are less accessible or endangered. Also, as goat production globally is vast and the market for goat products is expanding rapidly in the United States, studies of behavior and physiology of this species will contribute to enhanced productive performance and improved animal well-being. One application of this research may be to house males and females at a distance from one another if breeding is not intended, as the elevated androgenic hormones in males exposed to females can have negative impacts on growth and health.

Publications

• No publications reported this period

Progress 01/01/11 to 12/31/11

Outputs
OUTPUTS: Studies of female mate choice in goats were continued. Findings were presented at the annual meeting of the American Scoiety of Animal Science in New Orleans, LA. One journal article has been published so far and additional manuscripts are in preparation for peer review. PARTICIPANTS: Susan E. Becker, MS., was the research assistant participating in all phases of the experiments. Kristy Lonpre, PhD conducted all of the behavior experiments. She completed her PhD in 2011. In addition, dozens of undergraduate animal science students assisted with animal handling and data collection. TARGET AUDIENCES: Scientific community in the area of Animal Behavior, Reproduction, and Evolutionary Biology. PROJECT MODIFICATIONS: Nothing significant to report during this reporting period.

Impacts
One goal of this research was to demonstrate that the domesticated female goat (Capra hircus)may be used as a model for the study of female mate choice. Implications for this research may extend to a variety of other ungulate species that are less accessible or endangered. Also, as goat production globally is vast and the market for goat products is expanding rapidly in the United States, studies of behavior and physiology of this species will contribute to enhanced productive performance and improved animal well-being. One application of this research may be to house males and females at a distance from one another if breeding is not intended, as the elevated androgenic hormones in males exposed to females can have negative impacts on growth and health.

Publications

• Longpre K.M., Koepfinger, M.E. and L.S. Katz. 2011. Female goats use courtship display as an honest indicator of male quality. Hormones and Behavior 60: 505-511. 10.1016/j.yhbeh.2011.07.019
• Haulenbeek, A.M. and L.S. Katz. 2011. Female tail wagging enhances sexual performance in male goats. Hormones and Behavior 60:244-247. 10.1016/j.yhbeh.2011.05.008
• Longpre K.M. and L.S. Katz. 2011. Estrous female goats use testosterone-dependent cues to assess mates. Hormones and Behavior 59: 98-104. doi:10.1016/j.yhbeh.2010.10.014

Progress 01/01/10 to 12/31/10

Outputs
OUTPUTS: A series of experiments were conducted to address questions in the area of sexual selection, with particular focus on female mate choice. PARTICIPANTS: Kristy Lonpre, PhD student conducted all of the behavior experiments. In addition, dozens of undergraduate animal science students assisted with animal handling and data collection. TARGET AUDIENCES: Scientific community in the area of Animal Behavior, Reproduction, and Evolutionary Biology. PROJECT MODIFICATIONS: Nothing significant to report during this reporting period.

Impacts
In a promiscuous species like the domestic goat (Capra hircus), in which maternal investment is greater than paternal investment, a female may mate selectively with a more-fit male to improve her reproductive fitness. Testosterone (T) controls a large suite of male-typical behaviors and morphological characteristics. High T concentrations may be energetically costly or even detrimental to survival,thus preventing lower quality males from falsely advertising their fitness. Three preference studies were conducted to examine if females use T-dependent cues to assess potential mates. For Experiment 1, females were given a choice between a pair of morphologically similar males, bucks (intact males) and stags (post-pubertally castrated males), during the breeding and non-breeding seasons. In both seasons, females preferred the bucks compared to stags. In Experiment 2, females were given a choice between bucks, stags and wethers (pre-pubertally castrated males) during the non-breeding season. For some comparisons, castrated males received 25mg testosterone propionate (TP) or were untreated. Females preferred TP-treated males compared to untreated males and showed no preference when given a choice between either two TP-treated or two untreated males. In Experiment 3, females were given a choice between a pair of bucks and a pair of stags treated with 25mg TP during monthly tests in the breeding season. At each monthly test, females preferred the males with higher T concentrations near the time of the behavior test. These studies suggest that females use T-dependent cues to assess potential mates, and T concentrations may indicate a male's overall fitness. Sexual selection may have great influence on the life-history traits of males as reproductive effort, including the increased expression of secondary sexual characteristics and behaviors, improves a male's reproductive success. Alteration in T production depending on male quality and reproductive opportunities may provide males with the ability to invest differentially in reproductive effort to minimize reproductive costs. To test the effect of the presence of females on circulating T concentrations, males were housed with both fence-line and visual contact with females (Near bucks, n=5), or no fence-line nor visual contact with females (Far bucks, n=5), beginning Aug 4, prior to the onset of the breeding season. After 10 wk, the Far bucks were added to the Near so all had fence-line contact with females. Weekly blood samples were collected for the duration of the experiment. Far bucks had significantly lower T concentrations than Near for the first 10 wk. After fence-line contact with females, T concentrations for the Far bucks increased and were similar to those of the Near bucks. In addition, for both the bucks and TP-treated males, body mass loss was greatest when T concentrations were high. Taken together, these studies suggest that T may serve as a physiological mechanism that mediates life-history trade-offs for males.

Publications

• Longpre K.M. and L.S. Katz. 2010. Estrous female goats use testosterone-dependent cues to assess mates. Hormones and Behavior. doi:10.1016/j.yhbeh.2010.10.014
• Margiasso, M.E. Longpre, K.M. and L.S. Katz. 2010. Partner Preference: Assessing the Role of the Female Goat. Physiology and Behavior 99:587-591. doi:10.1016/j.physbeh.2010.01.022

Progress 01/01/09 to 12/31/09

Outputs
OUTPUTS: A series of experiments were conducted to address questions in the area of sexual selection, with particular focus on female mate choice. PARTICIPANTS: Kristy Lonpre, PhD student conducted all of the behavior experiments. In addition, dozens of undergraduate animal science students assisted with animal handling and data collection. TARGET AUDIENCES: Scientific community in the area of Animal Behavior, and Evolutionary Biology. PROJECT MODIFICATIONS: Nothing significant to report during this reporting period.

Impacts
In a promiscuous species such as the goat, Capra hircus, in which maternal investment is greater than paternal investment, a female may mate selectively with a more fit male, resulting in improved reproductive fitness. Previous research from our laboratory indicates that estrous females prefer males with higher serum testosterone (T) concentrations over males with lower concentrations, suggesting that females are using a T-dependent cue to assess potential mates. Further, our data provide evidence that both courtship display and chemical signals play an important role in the female's preference for a particular male. When estrous females were provided a choice between rags saturated with odors from bucks (gonad-intact males) versus stags (castrated males), they spent more time near the buck-scented rags. Estrous females also preferred bucks who courted them at a high frequency (120 courts/10m) compared to those who courted them at a low frequency (40 courts/10min). In two preliminary studies examining chemical signaling and serum T concentrations, we found that males display an increase in self-scent marking (self-enurination) early in the breeding season when T concentrations are high. This behavior decreased as the breeding season progressed and T concentrations normally decreased. In the second preliminary study we found that high-courting bucks have higher serum T concentrations compared to low-courting bucks, and bucks that courted at a medium frequency had serum T concentrations that fell between the high- and low-courting males. Taken together, these results suggest that both courting frequency and chemical cues provided by bucks may be honest indicators of T concentrations and thus buck quality. Additional studies are underway to further examine the relationship between courtship frequency, chemical cues and serum T concentrations.

Publications

• Longpre, K.M. and L.S. Katz. 2009. Females use testosterone regulated cues to assess potential mates. Society for Behavioral Neuroendocrinology, East Lansing, MI.

Progress 01/01/08 to 12/31/08

Outputs
OUTPUTS: Studies were conducted to address sexual partner preference in female goats, as well as studies of sexual stimulation in male goats. Presentations were made at the annual meeting of the Animal Behavior Society. The research was conducted by two graduate students as part of their doctoral training in Endocrinology and Animal Biosciences. In addition, approximately 40 undergraduate students were trained in animal behavior research as part of the experience-based educational requirement for the B.S. in Animal Sciences. PARTICIPANTS: Andrea Haulenbeek, PhD student. Kristy Longpre, PhD student Susan Becker, Research Assistant TARGET AUDIENCES: Scientific community PROJECT MODIFICATIONS: Nothing significant to report during this reporting period.

Impacts
New information was generated through the research. In the nonbreeding season, females (does) prefer testosterone (T)-treated stags (males castrated postpubertally) to bucks. In this study, in the peak of the breeding season, individual females were placed in a 17 m arena that had two bucks in a pen at one end and two T-treated stags in a pen at the other. Estrous does (n=11) spent more time near the bucks (P<0.001). Six weeks later, after stags had fence line contact with does, this preference was reversed (n=9). Serum T concentrations were higher for bucks than for stags near the time of each preference test. Prolonged T-treatment and/or close proximity to does may have increased sebaceous gland output in stags. During the same season, to test if olfactory cues influence female choice, the same arena had two wethers (males castrated prepubertally) at each end. The pens differed in scent cues by the addition of buck-scented rags to one and wether-scented rags to the other. Does (n=20) spent more time near the buck-scented rags (P<0.001), confirming that male-derived odors attract females. We propose the existence of a "female effect," in which contact with females increases male sebaceous gland output and sexual behavior in goats. We hypothesized that exposure to females expressing high rates of tail wagging would arouse males, increasing sexual performance. We have shown that flutamide treatment increases the frequency of tail wagging in estrous goats. Sexually-experienced males observed different stimuli for 10 min before a 20 min sexual performance test (SPT). The stimuli were an empty pen, or groups of three females that were all estrous (E), non-estrous (NE), estrous+flutamide (Ef) or non-estrous+flutamide (NEf). During the stimulus observation period, tail wagging was recorded. During SPT, frequencies and latencies of sexual behaviors were recorded. Ef females displayed the most tail wagging. Viewing Ef females before SPT significantly increased the number of ejaculations attained by males (2.9 vs. 1.6 for Ef and E, respectively). We conclude that male goats are sexually aroused by tail wagging. This study and previous work demonstrate that tail wagging functions as both attractivity and proceptivity in goats.

Publications

• Katz, L. S. 2008. Variation in male sexual behavior. Animal Reproduction Science 105: 64-71.
• Haulenbeek, A. M. and L.S. Katz. 2008. Female tail wagging enhances sexual performance in male goats, Capra hircus. Animal Behavior Society, Snowbird, Utah.
• Longpre, K.M., Becker, S.E. and L.S. Katz. 2008. Female mate choice is influenced by male olfactory cues in the domestic goat, Capra hircus. Animal Behavior Society, Snowbird, Utah.

Progress 01/01/07 to 12/31/07

Outputs
OUTPUTS: The research contributed to the thesis of two PhD candidates, as well as the work of a post-doctoral associate. Findings were presented at the annual meeting of the Society for Behavioral Neuroendocrinology during the summer of 2007. In addition, an invited presentation was made at a symposium at Colorado State University during January 2007. PARTICIPANTS: D. Bradley Imwalle, PhD.: Research associate. Andrea Haulenbeek: PhD student Kristy Longpre: PhD student Susan Becker, M.S.: Research assistant TARGET AUDIENCES: The target audience for this basic research are behavioral neuroendocrinologists, reproductive biologists, and animal scientists.

Impacts
Partner preference testing was used in which males could choose between two females located in pens at opposite ends of a test arena. Females were non-estrous, estrous or estrous and treated with flutamide. In previous work flutamide enhanced tail-wagging in estradiol-treated females. Thirteen males were tested in six 5 min trials. Time with and visits to each female were recorded. Bouts and duration of tail-wagging were recorded. Flutamide-treated females spent more time tail-wagging and displayed more bouts of tail wagging than other females. Males showed no preference between estrous and non-estrous females and between estrous and flutamide-treated females, but visited the flutamide-treated females the most. This indicates that the incentive properties of tail-wagging initiate approach behaviors, suggesting that tail-wagging is an indicator of attractivity. Males preferred flutamide-treated females over non-estrous females, visited flutamide-treated females most, further supporting that tail-wagging is an attractive behavior. Males also tended to spend more time per visit with flutamide-treated females, suggesting that tail-wagging may also be proceptivity, because after initial approach behaviors females continue to display the behavior and males maintain interest in the females. Males preferred females that tail-wagged most frequently and for the longest duration. Tail-wagging is an important attractive and proceptive behavior affecting partner preference and sexual motivation in the male goat. The promiscuous nature of mating in goats, along with greater reproductive investment by females, supports the hypothesis that females choose among males for a mate. To determine the cues that influence female partner preference, an arena (17m x 3.6m) was designed with a pen (1.5m x 3m) on each end to confine males. Dividing the length of the arena at the middle were two 2.4m wide pieces of 80% shade cloth, set 2.4m apart but overlapping to create 1.2m wide openings at each side. This visual barrier defined the neutral zone where estrous-synchronized females (n=12) were released individually and allowed 5 min to choose between two bucks in one pen or two stags (post-pubertal castrates) in the other. The area between the neutral zone and a pen was defined as an incentive zone (IZ; 5.9m x 3.6m). Females spent more time in the buck IZ (P<0.001), preferring the bucks to the stags, which looked like bucks but did not behave or smell like them. To examine if male odors influence female partner preference in the absence of buck-like visual cues, a second study was conducted. Buck-scented rags (BR) were tied to one pen in the arena, and control rags to the other. BR were strips of fabric that had been in the bucks' home pen for two weeks. Control rags were not exposed to goats. In addition, each pen contained two wethers (pre-pubertal castrates). Estrous-synchronized females (n=20) were tested individually for 5 min. Females spent more time in the BR IZ (P<0.001). These results show that estrous goats use olfactory cues when discriminating among males.

Publications

• Haulenbeek, A.M. and Katz, L.S. 2007. Female tail wagging elicits partner preference in male goats. Society for Behavioral Neuroendocrinology, Asilomar, CA.
• Longpre, K.M., Becker, S.E. and L.S. Katz. 2007. Estrous goats may use olfactory cues to choose prospective mates. Society for Behavioral Neuroendocrinology, Asilomar, CA.
• Imwalle, D.B., A. R. Lehrer and L.S. Katz. 2007. Intravaginal impedance and sexual behavior of ovariectomized goats given estrogen alone or in combination with progesterone. J. Animal Science 85: 1908-1913.
• Katz, L.S. 2007. Sexual behavior of domesticated ruminants. Hormones and Behavior 52: 56-63.
• Imwalle, D. B. and Katz, L.S. 2007. Suppression of the hypothalamic-pituitary-adrenal axis fails to affect sexual behavior in estradiol-treated ovariectomized goats. Society for Behavioral Neuroendocrinology, Asilomar, CA.

Progress 01/01/06 to 12/31/06

Outputs
Androgen receptor (AR) antagonism impairs receptivity in testosterone-treated, ovariectomized (ovx) goats, whereas it facilitates proceptivity in estradiol-treated, ovx goats. Androgens also have direct effects on intravaginal impedance (IVI). To test the effects of an AR antagonist on sexual behavior and IVI in gonad-intact goats, 12 sexually-experienced females were randomly assigned to receive flutamide (9 mg/kg) or vehicle. Intra-vaginal impedance was measured at 1 kHz (tissue extracellular space) and 70 kHz (combined extra- and intracellular spaces) using 1.5 mA current on d 1-6. Estrus was synchronized using injections of prostaglandin on d -9 and 3. Flutamide or vehicle was injected s.c. in the evening of d 2, the morning of d 3 and the evening of d 3. On d 5 cohorts of three females from the same treatment group were released into a yard containing a confined male (1.7 m x 3.2 m pen) for 10 min and the following variables were recorded: time spent within one body length of the male; time spent with head in the pen; frequency of head placement in the pen; frequency of female-female mounts. Females were also individually tested for 15 min for expression of sexual behaviors with a male. Proceptivity did not differ between treatment groups and all females were receptive. Flutamide-treated females had a significantly higher IVI value at the 1 kHz frequency on d 6 compared with vehicle-treated females. These results suggest that the AR mediates changes in the reproductive tract as measured by IVI, but is not essential for expression of sexual behavior in the gonad-intact female goat. Sexual motivation, the desire to engage in sexual activity, can be stimulated by the presence of suitable mates. Partner preference may be an indirect measure of sexual motivation. To test partner preference, a male goat in a rectangular pen (10 m x 4 m) could choose between an estrous female and an anestrous female, each in a pen (3 m x 2 m) at opposite ends of the rectangle. A 1.2 m incentive zone (IZ) was defined in front of each female's pen, with a 4 m neutral zone located between IZs. On four separate days, seven sexually-experienced males were tested once each d in 10 min periods for partner preference. Locations of the females were alternated to account for possible sidedness of the males. Time spent in each IZ and number of visits to each were recorded. In a fifth trial, the 10 min preference test was preceded by a 5 min sexual performance test (SPT). During the first four trials males spent more time in the estrous IZ (P<0.01). There was no difference in the number of visits to the IZs. After the SPT there was no longer a preference for estrous females as measured by time spent in IZ. In the first and third trials tail wagging frequency was low (but not quantified) in estrous females. Males preferred estrous females that tail wagged. We are currently studying the relationship between tail wagging in females and male partner preference. We conclude that the usefulness of partner preference tests to assess sexual motivation is influenced by the stimulus value of the choices.

Impacts
Major loses occur to animal agriculture due to inadequate expression of male and female sexual behaviors, which results in delayed reproduction and/or production of too few offspring. Research on the underlying physiological, endocrine, and environmental influences on reproductive behavior will contribute to the amelioration of these problems. These studies expand our understanding of androgen stimulation of reproductive behavior and corresponding changes in female reproductive tract function. In addition they are beginning to yield information that will contribute to the development of tests of sexual motivation as a potential predictor of sexual performance in males.

Publications

• Shearer, M.K. and L.S. Katz. 2006. Female-female mounting among goats stimulates sexual performance in males. Hormones and Behavior 50:33-37.
• Wohlt, J.E., Imwalle, D.B. and Katz, L.S. 2006. Use of recycled paper (news/office) and straw as bedding and their effects on heifer cleanliness and behavior. Federation of Animal Science Societies Joint Annual Meeting, Minneapolis, MN.
• Imwalle, D. B., Becker, S.E., and Katz, L.S. 2006. Novel approaches to teaching behavior to animal science students. Federation of Animal Science Societies Joint Annual Meeting, Minneapolis, MN.
• Haulenbeek, A. M. and L.S. Katz. 2006. Partner preference in male goats to measure sexual motivation. Society for Behavioral Neuroendocrinology, Pittsburgh, PA.
• Nichols, M. and L.S. Katz. 2006. An optimal dose of testosterone for activating sexual behaviors in ovariectomized goats. Society for Behavioral Neuroendocrinology, Pittsburgh, PA.
• Imwalle, D. B., Lehrer, A.R., and Katz, L.S. 2006. Androgen receptor antagonism, intravaginal impedance and sexual behavior in female goats. Society for Behavioral Neuroendocrinology, Pittsburgh, PA.

Progress 01/01/05 to 12/31/05

Outputs
1. Androgen receptor (AR) antagonism impairs receptivity in testosterone-treated, ovariectomized goats, whereas it facilitates proceptivity in estradiol-treated, ovariectomized goats. Androgens also have direct effects on intravaginal impedance (IVI). To test the effects of an AR antagonist on sexual behavior and IVI in gonad-intact goats, 12 sexually-experienced females were randomly assigned to receive flutamide (9 mg/kg) or vehicle. Intra-vaginal impedance was measured at 1 kHz (tissue extracellular space) and 70 kHz (combined extra- and intracellular spaces) using 1.5 mA current on Days 1-6. Estrus was synchronized using injections of prostaglandin on Days -9 and 3. Flutamide or vehicle was injected s.c. in the evening of Day 2, the morning of Day 3 and the evening of Day 3. On Day 5 cohorts of three females from the same treatment group were released into a yard containing a confined male (1.7 m x 3.2 m pen) for 10 minutes and socio-sexual behaviors were recorded. Females were also individually tested for 15 minutes for expression of sexual behaviors with a male. Proceptivity did not differ between treatment groups and all females were receptive. Flutamide-treated females had a significantly higher IVI value at the 1 kHz frequency on Day 6 compared with vehicle-treated females. It was concluded that the AR mediates changes in the reproductive tract as measured by IVI, but is not essential for expression of sexual behavior. 2. Sexual motivation, the desire to engage in sexual activity, can be stimulated by the presence of suitable mates. Partner preference may be an indirect measure of sexual motivation. To test partner preference, a male goat could choose between an estrous female and an anestrous female, at opposite ends of a large pen. On four separate days, seven sexually-experienced males were tested once each day in 10 min periods for partner preference. Regardless of estrous state, males preferred estrous females that tail-wagged. We concluded that the usefulness of partner preference tests to assess sexual motivation is influenced by the stimulus value of the choices. 3. We are developing a model to study environmental and social factors that influence androgen-mediated sexual behavior in the female goat. This necessitates determining an effective dose of testosterone (T) just above the minimum threshold for activating estrous behaviors (optimal dose). The optimal dose of T required for inducing proceptivity, attractivity and receptivity in ovariectomized (ovx) French-Alpine goats was investigated during the breeding season. For goats receiving 500 ug T or more, proceptivity and attractivity scores were each greater than for controls (P<0.05). Receptivity scores were significantly higher only following administration of 750 and 1000 ug, however the proportion of females displaying receptivity increased following administration of 250 ug (50%) compared to controls (0%; P<0.02). Furthermore, following treatment with 500 ug T 80% of the females were receptive. Taken together these data suggest that 500 ug is the optimal dose of T for studying the induction of sexual behavior in ovx female goats.

Impacts
Major loses occur to animal agriculture due to inadequate expression of male and female sexual behaviors, which results in delayed reproduction and/or production of too few offspring. Research on the underlying physiological, endocrine, and environmental influences on reproductive behavior will contribute to the amelioration of these problems. These studies expand our understanding of androgen stimulation of reproductive behavior and corresponding changes in female reproductive tract function. In addition they are beginning to yield information that will contribute to the development of tests of sexual motivation as a potential predictor of sexual performance in males.

Publications

• Margiasso, M.E. and L.S. Katz. 2005. Use of a Y-Maze to Test Partner Preference in female Goats. Society for the Study of Reproduction, Quebec, Canada.
• Sweeney, A. and L.S. Katz. 2005. Use of a Y-Maze to Test Partner Preference in Male Goats. Society for the Study of Reproduction, Quebec, Canada.
• Imwalle, D.B., Lehrer, A.R. and L.S. Katz. 2005. Estradiol initiates and progesterone sustains the drop in vaginal impedance at estrus in ovariectomized goats. Society for the Study of Reproduction, Quebec, Canada.

Progress 01/01/04 to 12/31/04

Outputs
Female goats in estrus mount one another. We have previously shown that the frequency of this behavior increases in the absence of male goats or when male goats are inaccessible to the females. The effects of watching female-female mounting on male sexual performance have not been described. We hypothesized that female-female mounting stimulates the sexual performance of males, and therefore can be considered proceptive behavior in this species. A randomly selected male (n=6) was placed in a test pen and given the opportunity to observe one of six randomly selected stimuli in an adjacent pen. The stimuli were: an empty pen; three familiar males; three non-estrous females; three estrous females that displayed female-female mounting; three estrous females that did not display female-female mounting; and one male with two estrous females. After 10 minutes the stimulus animals were removed and an estrous female was introduced into the test pen with the male for 20 minutes, during which, the frequencies and latencies of all sexual behaviors were recorded. This procedure was repeated for each of the remaining males, such that all males were tested once and all of the stimulus conditions were presented on any given day. The experiment was repeated weekly, each male being tested under each condition in random order. Providing male goats the opportunity to view mounting behavior, whether male-female or female-female, increased total number of sexual behaviors displayed, increased ejaculation frequency, and decreased latency to ejaculation, post-ejaculatory interval and the interval between ejaculations. We conclude that male goats are aroused by visual cues from other mounting animals and that female-female mounting is a proceptive behavior.

Impacts
This study provides further evidence that sexual interactions among individuals of the same gender, referred to as heterotypical behavior, functions as an important component of the socia-sexual behavior of animals. Also, the frequent occurrence of heterotypical sexual behavior among goats suuports the use of this species to better understand normal as well as dysfunctional sexual behavior in humans.

Publications

• Imwalle, D.B. and L.S. Katz. 2004. Divergent roles for estrogens and androgens in expression of female goat sexual behavior. Hormones and Behavior 46: 54-58.
• Imwalle, D.B. and L.S. Katz. 2004. Development of sexual behavior over several serving capacity tests in male goats. Applied Animal Behaviour Science 89:315-319.
• Katz, Larry S. Behavior: Reproductive. 2004. In Encyclopedia of Animal Science; Pond, Wilson G., Bell, Alan W., Eds.; Marcel Dekker, Inc.: New York.
• Imwalle, D.B., and L.S. Katz. 2004. An aromatase inhibitor fails to suppress estrous behavior in testosterone-treated ovariectomized goats. Society for Behavioral Neuroendocrinology, Lisbon, Portugal, Hormones and Behavior.
• Imwalle, D.B., and L.S. Katz. 2004. Repeatability of serving capacity scores in male goats requires prior sexual experience. International Society of Applied Ethology-North American Region, Purdue, IN.
• Sweeney, A., Imwalle, D.B. and L.S. Katz. 2004. Assessment of motivation in male goats as a predictive tool for sexual performance. International Society of Applied Ethology-North American Region, Purdue, IN.

Progress 01/01/03 to 12/31/03

Outputs
We tested the hypothesis that the activation of the androgen receptor (AR) is required for full expression of female goat sexual behavior. Once a week for 6 weeks, ovariectomized (OVX) females were given priming doses of progesterone 72 and 48 h prior to behavioral observation. Estradiol (E2; 100 ug), testosterone (T; 100 mg) or sesame oil was supplied 14 h prior to behavioral testing. Six goats received the AR antagonist flutamide (9 mg/kg sc) 8 h prior to and 4 h after steroid injection. Six goats received the carrier only. After 3 wk, flutamide and carrier treatments were switched so that all females received all treatments. Treatment with E2 and T were equally effective in eliciting estrus-typical behaviors (sniffing, courting, leg kicks, mount attempts by males, bouts of thrusting by males, ejaculations, and flehman responses) compared to treatment with oil. Flutamide treatment enhanced proceptive behaviors in E2-treated females compared to other treatment groups; this was most likely via enhanced tail wagging. Moreover, compared to goats given T+carrier, T+flutamide significantly reduced receptivity in females. The results of this experiment implicate the AR as an important facilitator of some aspects of female goat sexual behavior. However, the results of this experiment do not show whether androgens influence estrous behaviors alone on in some combination with estrogen.

Impacts
In sum the results of this experiment suggest divergent roles for estrogens and androgens as mediators of female goat sexual behavior. Whereas estrogens appear to be responsible for facilitating proceptivty, and possibly attractivity, receptivity in female goats appears to be influenced by an androgen-dependent pathway. However, the results of this experiment do not show whether androgens influence estrous behaviors alone on in some combination with estrogen. Nevertheless, these observations may make the goat a powerful comparative model for the human female, given the great interest in androgen-facilitation of libido in post-menopausal women

Publications

• Imwalle, D.B. and L.S. Katz. 2004 in press. Divergent roles for estrogens and androgens in expression of female goat sexual behavior. Hormones and Behavior.
• Imwalle, D.B., and L.S. Katz. 2003. The effects of flutamide on ovariectomized goat sexual behavior. Hormones and Behavior 44:56.
• Lindia, J.A., and L.S. Katz. 2003. Estradiol and testosterone stimulation of sexual behavior in ovariectomized female goats. Hormones and Behavior 44:62.
• Shearer, M., and L.S. Katz. 2003. Female-female mounting among goats stimulates sexual performance in males. Hormones and Behavior 44:77.

Progress 01/01/02 to 12/31/02

Outputs
We performed a series of experiments that investigate the possible roles of androgens in female sexual behavior. Female French-Alpine ovariectomized goats were treated in a Latin square design for each study. Progesterone (P4)-filled controlled intravaginal drug release devices (CIDRs) were inserted into the vagina and removed 72 hours later. Steroid treatments were given subcutaneous 24 hours after removal of CIDR. At 15 and 20 hours following treatment sexual behavior observations were conducted. A sexually experienced, testosterone-treated wether or ram was placed with a female for ten minutes. During that time, specified sexual behaviors were recorded. From the recorded behaviors, female attractivity and receptivity were determined. The first study compared the ability of estrogen (E2), testosterone (T), dihydrotestosterone (DHT) and androstenedione (A4) to stimulate estrous behavior. The second study, determined whether there is a dose response of E2 and T concentrations on the induction of estrous behavior. With the knowledge gained from the dose response, two additional studies were completed. In the first of these studies, we co-administered the non-aromatizable androgen, DHT, with a sub-threshold dose of E2. In the second study, we administered an aromatizable androgen, T, and E2 in an attempt to determine if combining two known inducers of estrous behavior would amplify their overall effects. From these studies, we learned that only aromatizable androgens stimulate estrous behavior. The data suggests the T and A4 may be more effective at bringing a goat into estrus than E2. The dose response studies suggest that there is no linear effect of dose versus estrous behavior when estradiol is given at various levels above the threshold dose. Furthermore, low dose testosterone is no more effective than a high dose. Dihydrotestosterone, given concurrently with a sub-threshold level of E2, cannot induce estrous behavior. Lastly, we determined that the effects of maximal E2 and T couldn't be increased even when given together. Other projects have included a descriptive analysis of the behavior of cloned cattle; and more recently we have demonstrated that the opportunity to observe female goats mounting each other improves sexual performance in male goats.

Impacts
The efficacy of androgens to stimulate sexual activity in females can be applied in estrous detection programs. Sexual performance of goats can be improved by providing visual contact with females. The behavioral potential and possible side-effects of medical application of androgens for post-menopausal women can be studied using goats.

Publications

• SAVAGE, A.F., MAULL J., TIAN X.C., TANEJA M., KATZ L., DARRE M., and YANG X. 2003. Behavioral Observations of Adolescent Holstein Heifers Cloned from Adult Somatic Cells. Theriogenology.

Progress 01/01/01 to 12/31/01

Outputs
No progress to report -- too early in the life of the project to report any activities.

Impacts
(N/A)

Publications

• No publications reported this period