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ENVIRONMENT, WELL-BEING, AND BEHAVIOR |
Applied Animal Biotechnology Laboratories, School of Animal Sciences, Louisiana Agricultural Experiment Station, Louisiana State University Agricultural Center, Louisiana State University, Baton Rouge 70803
2 Corresponding author: dsatterlee{at}agctr.lsu.edu
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ABSTRACT |
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 TOP ABSTRACT INTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONREFERENCES |
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Key Words: Japanese quail • tonic immobility • timidity • corticosterone • stress
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INTRODUCTION |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONREFERENCES |
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It now appears that plasma B elevations in hens during egg formation (Hayward and Wingfield, 2004), or in ovo B-treatment (Rubolini et al., 2005; Janczak et al., 2006), can also enhance stressor-induced sensitivity of the hypothalamic-pituitary-adrenal (HPA) axis of the young of the hen, which further translates into negative consequences on the performance and behavior of the offspring. Specifically, when hens were implanted with B during egg formation, more B was deposited into the egg yolk and chicks of B-implanted mothers showed reduced growth and a higher HPA activity in response to restraint as adults (Hayward and Wingfield, 2004). Chicks hatched from in ovo B treatments also show a reduced food drive and more fear of humans (Janczak et al., 2006). Additionally, B-treated yellow-legged gull hen eggs produce chicks with decreased cell-mediated immunity, a reduced rate and loudness of late embryonic vocalizations, and attenuated intensities of chick begging (Rubolini et al., 2005). Mice offspring from mothers experiencing harsh (i.e., stressful) prenatal conditions also show less environmental exploration (Benderlioglu et al., 2006).
As reviewed above, when compared with LS quail, HS quail show an exaggerated plasma B response to many different nonspecific systemic stressors, as well as heightened fearfulness. Because Hayward and Wingfield (2004) found genetically unremarkable quail hens given B to deposit more B in the yolks of their eggs and produce adult offspring with an exaggerated HPA responsiveness to brief restraint (a trait shared with our HS quail), because Janczak et al. (2006) found in ovo B-treatment to increase fear in hatchling chicks, and because Hayward et al. (2005) also found that HS hens deposit more B into their egg yolks than do LS hens, we wondered whether maternal B treatment would interact with the LS and HS quail genomes to alter fear responses in their offspring. The present study tested this hypothesis. Specifically, during egg formation, LS and HS mothers were given implants filled with either B or no B (controls), and then their juvenile offspring were tested for differences in underlying fearfulness (via tonic immobility, TI, tests) and in the timidity aspects of fear (using hole-in-the-wall box, HWB, emergence testing).
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MATERIALS AND METHODS |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONREFERENCES |
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Offspring from generation 38 of 2 lines selected for either low (LS) or high (HS) plasma B response to brief immobilization were studied. Satterlee and Johnson (1988) have described the genetics that underlie the first 12 generations of pedigree selection and the most recent genetic history of the lines, up to generation 34, is discussed in detail elsewhere (Satterlee et al., 2000, 2006; Marin and Satterlee, 2004). Although line differences in levels of plasma B were not measured in the present study, recent findings in the stress lines attest to the maintenance of divergent adrenocortical responsiveness to a variety of nonspecific systemic stressors. Indeed, Satterlee et al. (2007) have most recently offered explanations as to why we believe that the gene(s) that control the adrenocortical responsiveness trait in these lines have become fixed.
Ninety-six hens from each line (48 LS + 48 HS) were used. At 29 wk of age, each hen was pair-housed with a nonsibling, same-line male in a single cage of 1 of 2 Alternative Cage Designs (Alternative Design Manufacturing and Supply Inc., Siloam Springs, AR) 4-tier cage batteries. Each battery contained 48 pedigree-style breeder cages (individual cage dimensions were 50.8 x 15.2 x 26.7 cm, length x width x height, respectively). Care was taken to ensure that each of the breeding pairs selected, while randomly selected from larger family populations within each line of the same hatch, constituted, as nearly as possible, equal representation of the 12 different families that make up each line. A breeder ration (21% CP; 2,750 kcal of ME/kg) and water were provided to the birds ad libitum. The daily photostimulatory cycle was 14L:10D (approximately 280 lx during the lighted portion of the day); lights-on was at 0600 h and lights-off was at 2000 h daily. Daily maintenance and feeding chores were done at 0800 h daily.
Hen Treatments
At 33 wk of age, half of the hens from each line (n = 24 birds/line) were individually fitted with 16-mm silastic tube (Dow Corning, Midland, MI; cat. no. 508-006) implants containing either corticosterone (B; Sigma-Aldrich Co., Atlanta, GA; cat. no. C2505) or no B (controls, CON). Implants were placed s.c. in the back of the neck using a No. 10 biopsy needle (Becton Dickinson, Franklin Lakes, NJ). The implant tubes were sealed at one end with silicone sealant and open on the other end. Hens were allowed a 10-d acclimation period to allow sufficient time for maternal B deposition into the eggs of B-treated hens (Hayward and Wingfield, 2004) to their implantation treatments. Eggs were then collected daily, identified by pencil markings as to their origin by hen line and implantation treatment, and stored at 18°C until incubation. Egg collection lasted for 3 wk, and these eggs were then set together into an incubator (NatureForm NMC 2000; NatureForm Hatchery Systems, Jacksonville, FL). During the first 14 d of incubation, eggs were turned 6 times a day and subjected to 37.5°C and 62% RH. Upon transfer of the eggs to a second NMC 2000 hatcher unit on d 14, eggs were no longer turned, and incubation conditions were changed to 37.2°C and 69% RH.
Offspring and Variables Measured
At hatch, chicks were leg-banded with appropriate different color and uniquely numbered leg bands that allowed their identification with the 4 line x implantation treatments (LS-CON, LS-B-implant, HS-CON, and HS-B-implant). Chicks were brooded, and all treatments were equally comingled, in 3 confinement rings (approximately 260 chicks/ring). This arrangement resulted in about 65 chicks from each treatment combination being represented in each ring. The brooding ring areas were of identical construction: each ring was 1.2 m in diameter, heated with two 125-W incandescent lamps, and had pine wood shavings as a floor substrate. Chicks were fed a quail starter ration (28% CP; 2,800 kcal of ME/kg) and given water ad libitum. Brooding temperatures and their change with time were similar to those used by Jones and Satterlee (1996).
At 14 d of age, 80 chicks (20 from each of the 4 stress line x implantation treatment combinations) were selected for TI studies. Individuals were randomly captured throughout the test day in equal rotation from each of the 3 confinement rings until the above sample numbers were achieved. Upon capture, a test chick was removed to a separate room (i.e., the TI test apparatus was located in a quiet area, approximately 13 m away from the live-bird facility, and free from bird noises and human traffic) and its TI responses were measured as follows. Placement of a bird on its back in a 120° V-shaped metal cradle covered by a white cloth was used to induce TI. Chicks were restrained in this dorsal recumbent position for 15 s using one of the hands of the experimenter placed on the sternum and the other lightly cupping the head. Successful induction into TI was defined as when a chick attained TI lasting for at least 10 s. If a chick did not achieve TI on the first try, additional induction attempts were made. If induction into TI was not accomplished after 5 attempts, a test subject was deemed unsusceptible to TI and given a score of 5 for the number of induction attempts (INDS) needed to induce TI. After a successful induction into TI, the experimenter quietly retreated to a nonintrusive position (approximately 2 m away from the TI cradle) while remaining in full sight of the chick. The experimenter then observed and recorded the latency from the end of induction into TI until the first alert head movement (generally a gross scanning behavior; LATHEAD, s) and the duration of TI (the length of time between the end of induction to observation of a chick self-righting response (TI, s). Maximum scores of 600 s (using a test ceiling of 10 min) were assigned to birds that showed no head movements (LATHEAD) and no self-righting behavior (TI) by the end of the test period. To ensure the continued capturing of untested chicks from their brooding environment, tested chicks were housed elsewhere.
The above experimental procedures were duplicated at 15 d of age, which served as an experimental replication (i.e., an additional 80 untested chicks, 20 birds per stress line x implantation treatment were TI tested). To minimize separation distress during testing on each day of the study, approximately only 10% of the commingled representatives of each treatment combination housed in a brooding area were tested daily.
At 21 d of age, an additional 80 chicks (20 each from the remaining untested LS-CON, LS-B, HS-CON, and HS-B treatment groups) were randomly selected for HWB studies. Individuals were again randomly captured throughout the test day in equal rotation from each of the 3 confinement rings until the above sample numbers were achieved. Upon capture, a test chick was removed to the same quiet room used for TI tests (see above) and its HWB responses were measured as follows. The testing box had 2 compartments (1 dark and 1 lighted) measuring 21 x 21 x 21 cm (length x width x height). The dark compartment was constructed of aluminum with a 1-cm wire mesh floor, and the lighted compartment was made entirely of wire mesh. Separating the 2 compartments was an aluminum wall with a 10 x 8-cm hole (height x width) covered by a guillotine trap door. The birds were placed individually into the dark compartment and given a 1-min acclimation period after which the guillotine door was raised.
The number of chicks that vocalized as a proportion of the total number of chicks tested (PVOCS), the latency to first vocalize (LATVOC, s), and the number of vocalizations (VOCS) before the guillotine door was raised were recorded. Chicks that did not vocalize during the 60-s acclimation period were given scores of 60 for LATVOC and 0 for VOCS. The number of chicks that vocalized during the 1-min acclimation period in the dark box as a proportion of total number of chicks tested in a line x implantation treatment combination (PVOCS) were also determined. The latencies from raising the door to head emergence through the hole in the wall of the dark box (head emergence, HE, s) and complete body emergence into the lighted compartment (full emergence, FE, s) were also recorded. Using a test ceiling of 10 min, maximum scores of 600 s were given to a chick that did not exhibit HE or FE behavior. Tested birds were rehoused in an area separate from their home-brooding area to ensure the capture and HWB testing of only untested chicks.
An experimental replication was performed at 23 d of age (i.e., the HWB responses of an additional 80 untested chicks, 20 birds per stress line x implantation treatment were determined). To minimize separation distress during HWB testing on each day of the study, approximately only 10% of the commingled representatives of each treatment combination housed in a brooding area were tested daily.
Statistical Analyses
The INDS, LATHEAD, TI, LATVOC, VOCS, HE, and FE data were subjected to nonparametric randomized block ANOVA that incorporated 2 x 2 factorial arrangements of treatments. The factorial was made on the effects of stress line (LS vs. HS) and maternal implantation treatment (CON vs B-implant). The blocks or experimental replications were made on the 2 consecutive days of observation in each experiment (14 and 15 d of age for TI tests; 21 and 23 d of age for HWB emergence tests). Duncan’s test was used to partition line x implantation treatment interaction differences in mean IND, LATHEAD, TI, LATVOC, VOCS, HE, and FE responses. The PVOCS variable is a binary trait (i.e., chicks either vocalized or not); therefore, a standard proportion test of differences was used for this variable.
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RESULTS |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONREFERENCES |
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The HS chicks required less (P < 0.0005) INDS to achieve TI than did the LS ones (Figure 1
, top panel; main effect mean, MEM). However, maternal implantation treatment did not alter INDS (Figure 1, middle panel; MEM), and post-hoc partitioning of the line x implantation treatment interactive effects (Figure 1, bottom panel) showed that both HS-CON and HS-B-implant treatments required similar and lesser (P < 0.01) numbers of INDS than did either of the 2 similarly responding LS treatments. On average, the LS chicks also took less (P < 0.02) time to show their first alert head movement (LATHEAD) after successful induction into TI than did the HS chicks (Figure 2, top panel; MEM), but neither the effect of maternal implantation treatment (Figure 2, middle panel; MEM) nor its inter-action with stress line (Figure 2, bottom panel) were effective in altering LATHEAD. Mean durations of TI were unaffected by line, implantation treatment, or their interaction (Figure 3).
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During the acclimation period in the HWB dark compartment, on average, the number of chicks that vocalized as a proportion of the total number tested (PVOCS) was much greater (P < 0.0001) for chicks of the HS line (Figure 4, top panel; MEM). The HS chicks also vocalized much sooner (P < 0.0001; LATVOC, Figure 5, top panel; MEM) and much more often (P < 0.0001; VOCS, Figure 6, top panel; MEM) than did the LS chicks. And, although maternal implantation treatment did not affect PVOCS (Figure 4, middle panel; MEM) or LATVOC (Figure 5, middle panel; MEM), CON chicks tended to vocalize more (P < 0.07) than did B-implanted ones (VOCS, Figure 6, middle panel; MEM). The stress line x implantation interactive effects on mean PVOCS and LATVOC were nonsignificant, and the post-hoc analyses of the mean responses of these effects (Figures 4 and 5, bottom panels, both P < 0.01) simply reflected the main effect of stress line (i.e., more HS chicks vocalized sooner than LS ones regardless of implantation treatment). However, a marginal (P < 0.07) line x implantation treatment interaction was found for VOCS. Post-hoc partitioning of the interactive VOCS means showed that the HS-CON chicks talked more (P < 0.01) than the other 3 similarly less talkative groups (Figure 6, bottom panel).
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and 8, respectively; MEM) into the lighted compartment of the HWB, mean HE (Figure 7, middle panel; MEM) and FE (Figure 8, middle panel; MEM) occurred sooner (P < 0.06 and P < 0.05, respectively) in the CON than in chicks hatched from B-implanted hens. The lines and implantation treatments showed no interaction in affecting HE and FE (bottom panels of Figures 7 and 8, respectively).
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DISCUSSION |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONREFERENCES |
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Herein, we again (Jones et al., 1992b; Satterlee et al., 1993) found LS chicks to show shorter LATHEAD behavior once inducted into TI than HS chicks. Because one of the first actions a bird may take before righting itself from TI is that of an alert lifting of its head (Jones, 1986), it follows that the novel TI test conditions or the experimenter, or both, are likely being perceived as less of a threat by LS than HS chicks. The idea that LS chicks may perceive humans as less of a threat has been proposed by Jones et al. (1994), who demonstrated that LS chicks show less avoidance of the faces of both a familiar caretaker and an unfamiliar experimenter than do HS chicks. Furthermore, when compared with HS quail, LS quail also show less avoidance of a novel object, such as a multicolored fishing float placed in their feed troughs (D. G. Satterlee, Louisiana State University, and R. B. Jones, Roslin Institute, unpublished data). Consistent with the antipredator (fear) TI hypothesis, avians consider natural and simulated natural predators (e.g., stuffed hawks), humans, artificial eyes, and even their own mirror reflections as predators in exhibiting TI behavior (Gallup, 1977).
Because of the line differences found in the INDS (LS > HS) and LATHEAD (LS < HS) behaviors, the duration of TI was expected to be reduced in LS quail as well. However, although HS quail remained in TI on average about 16% longer than LS quail, this difference was only numerically and not statistically relevant. The present lack of a significant line difference in the duration of TI contrasts with our previous studies that found a longer duration of TI in HS than LS quail of a similar age (Jones et al., 1992b; Satterlee et al., 1993). This discrepancy cannot be readily explained. However, when measuring fearfulness, certain fear-related variable results may not be consistently found between studies that have B-treatments in common. Indeed, Cockrem’s (2007) review on the interrelationships between stress, B, and avian personalities concludes that "although activation of the HPA axis when an animal perceives a stimulus to be a threat is considered to occur simultaneously with the basic emotion of fear" it can be challenging to relate fearfulness to B responses, because oftentimes "individual measures [of fear behavior] are not sufficient to quantify fear in birds." In other words, fear may not always be accurately measured by assessment of a single fear-related variable. In fact, Cockrem (2007), in reviewing one of his own studies (Fraisse and Cockrem, 2006), wherein the duration of individual TI measures to handling-induced B responses in the same chickens could not be linked, found that, by combining observations from 4 components of TI testing (INDS, LATHEAD, numbers of alert head movements, and duration of TI) into a fear score rank index, a significant correlation between this index and B responses resulted. This helps explain why the relationship between adrenocortical activation and heightened fearfulness in our quail stress lines (Jones et al., 1988, 1992a, b, 1994, 1999; Satterlee et al., 1993; Jones and Satterlee, 1996) is not always straightforward. Nevertheless, the present TI findings (considering that 2 of the 3 TI behavior variables measured showed line differences) do little to change the original contention (Jones, 1987, 1996) that fear and distress (adrenocortical activation) are positively correlated.
New to the present experiment is the assessment of influences of B-implant treatments in LS and HS hens during egg formation on their offspring’s TI behaviors. Surprisingly, neither maternal B-implantation treatment nor its interaction with stress line affected INDS, LATHEAD, or the duration of TI. These findings indicate that exogenous B treatment of stress line hens does not further alter the line differences in fear (HS > LS) detected in our previous studies (Jones et al., 1992b; Satterlee et al., 1993) and herein. These results were unexpected for the following reasons. First, B-implant treatment in genetically unremarkable quail hens (Hayward and Wingfield, 2004), a treatment identical to that used herein, is known to increase in ovo levels of B and heighten HPA responsiveness to brief immobilization in adult offspring of B-treated mothers. Moreover, the restraint stressor used by Hayward and Wingfield (2004) is the same stressor used to genetically select our LS and HS lines. Thus, we felt that, when compared with progeny of LS-CON hens, the offspring of B-implanted LS mothers would be prime candidates for maternal B-induced conversion into birds that would express greater adrenal responsiveness to stress and therefore be more fearful, attributes present as a result of genetic selection in our HS quail. We further reasoned that B-treatment of HS hens may or may not further exacerbate the stress x fear relationship in their HS chicks, depending upon whether selection for exaggerated adrenocortical responsiveness in the HS line has been maximized or not. We felt our preexperimental expectations were also reasonable considering the report in chickens (Janczak et al., 2006) that fear (avoidance of humans) is increased in chicks hatched from eggs injected with B.
Thus, the question remains: how does one explain the present lack of maternal B-implant influences on TI behaviors? It may be that genetic selection for divergent B response to stress has altered the genomic controls of adrenocortical responsiveness of quail lines in such ways that the HPA axis and therefore fear activity of stress line progeny cannot be further altered by whatever yet unidentified mechanism(s) that additional in ovo B during embryogenesis apparently uses to alter HPA activity (Hayward and Wingfield, 2004) and fear behavior (Janczak et al., 2006) in unselected avians. That said, however, it is important to note that Janczak et al. (2007b) have also found that although Leghorn hens stressed by feed restriction secreted more fecal B metabolites, levels of B in the albumen and yolks of their eggs were unaffected, yet the adult progeny of these stressed hens still showed longer durations of TI. This suggests that altered fear responsiveness in the offspring of these stressed mothers was the result of some other mechanism(s) independent of in ovo B. In yet a third study, Janczak et al. (2007a) found in ovo B injections during embryogenesis ineffective in altering TI responses in 4-wk-old chicks hatched from the B-treated eggs. Rubolini et al. (2005) have also found no changes in TI responses of yellow-legged gull chicks hatched from B-injected eggs. Thus, the 3 Janczak chicken studies and the gull study of Rubolini et al. (2005) present an unclear picture of the relationship between maternal and in ovo B influences on altering the TI behavior of avian offspring. Like the literature-proposed relationship between increased adrenal stress responsiveness and heightened fear, the avian maternal (or in ovo) B x offspring fear relationship is also not straightforward, and it awaits further clarification.
Presently, during the acclimation period to the dark compartment of the HWB, PVOCS were dramatically higher in HS than LS quail, and HS chicks also showed markedly reduced LATVOC and greater VOCS than LS chicks. Line differences in the LATVOC and VOCS behaviors have not been assessed previously, but the present PVOC result contrasts with an earlier study wherein more LS chicks vocalized than did HS ones during HWB acclimation (Jones et al., 1999). In that study, we concluded that the line difference in the number of chicks that vocalized during HWB acclimation was indicative of and consistent with the lower fear reactions (i.e., sooner HE and FE from the HWB once the guillotine door was raised allowing access to a presumably more frightening unfamiliar lighted space) also observed in that study – findings all consistent with the reduced fear reactions of LS quail found in numerous forerunner studies using other measurements of fear (e.g., open field behaviors, Jones et al., 1992a; TI, Jones et al., 1992b, Satterlee et al., 1993; avoidance of humans, Jones et al., 1994; and struggling in a crush cage, Jones and Satterlee, 1996).
However, a silence x fear hypothesis in HWB testing has not been just one-sided. For example, Leghorns deemed to be docile (assumedly less fearful birds) had a shorter LATVOC in emergence testing than did their flighty counterparts (Jones and Mills, 1983). The authors also submit that most would agree it is at best difficult for humans to know exactly what birds are saying when they vocalize, and admittedly, the authors are not expert in interpreting quail speak. However, reviews by Jones (1987, 1996) may be helpful here. They suggest that fear can either elicit or inhibit different numbers and qualities of vocalizations in avians depending upon the degree of novelty of environmental stressors as they relate to levels of fear. For example, low levels of novelty and fear have been associated with peeping distress calls, whereas intermediate fear levels often induce high-intensity peeping, and high levels of fearfulness may suppress vocalizations. We have the distinct impression that the vocalizations previously measured in our stress lines (vocalizations that were soft, infrequent, and only seen in the few birds that vocalized; in fact, 59 of the 80 earlier tested quail did not vocalize at all; Jones et al., 1999) were different than those heard presently, which were of a very frequent, high-pitched, more harsh, and louder nature – more like alarm calling (see below). The lack of much calling and the gentle nature of the VOCS of the few birds that called in our previous study (Jones et al., 1999) suggests that the fear levels associated with HWB testing produced then were likely much greater than what occurred in the present study. This may help explain why the line differences in PVOC contrasted between our 1999 (LS > HS) and present (HS > LS) studies. Indeed, Boissy’s (1995) review concludes that many avian species (e.g., magpies and domestic fowl) vocalize in the form of what has been called alarm calls when they detect predators. He defines alarm calls as signals, sounds whose structure can be varied in a graded fashion that communicate possibilities of danger to conspecifics. Such calling is believed to be influenced by physical characteristics of a frightening stimulus that is perceived to be predatory –factors such as stimulus presentation, movement, intensity duration, suddenness, or proximity. For example, Evans and Marler (1992) found the velocity of flying hawks in video images and image manipulation of visual distances from this known chicken predator were crucial in eliciting alarm calls in test (prey) chickens. Boissy (1995) has also suggested that novelty (such as what bird capture by the experimenter, transport to the HWB apparatus, separation from live conspecifics, and placement into the dark compartment of the HWB would have represented to the test birds of the present experiment) is "one of the most potent experimental conditions" that can lead to negative emotional responses, such as alarm calling. Moreover, exposure of chicks to alarm calls recorded from conspecifics exposed to a predator made them peep and run away, whether they have previously encountered a predator or not (Duncan and Filshie, 1979). Therefore, it is possible the present increased PVOCS, decreased LATVOC, and increased VOCS in HS quail are a simple reflection of greater antipredatory behavior of heightened alarm calling (i.e., HS quail perceived the nature of the cascade of events from human bird capture to placement in the dark compartment of HWB to be a more predacious experience than did LS quail). It is unfortunate that the vocalizations of LS and HS quail were not recorded so that they could be used in audio playback studies that would assess differences in the observed line vocalizations on peeping and runaway behaviors of unselected quail.
It is also important to note here that since our earlier report (Jones et al., 1999), in ovo B-treatment was found to increase the number and intensity of VOCS in yellow-legged gull hatchlings (Rubolini et al., 2005) and the numbers of distress vocalizations in domestic chicks following release into a novel arena (Freire et al., 2006). In addition, Viérin and Bouissou (2003) have used the utterance of more frequent high-pitched bleats to judge levels of fear in lambs in distress. Ultrasonic and audible fear-induced alarm call responses have also been documented in rats (assumed to be distress calls related to anxiety; Kikusui et al., 2001, 2003), squirrel monkeys (McCowan et al., 2001), and ground squirrels (Wilson and Hare, 2004).
Neither B-implantation treatment nor its interaction with quail stress line affected the HWB vocalization parameters measured herein. These results indicate B-treatment of LS or HS hens does not alter the vocalization activity of their respective offspring beyond the effects of stress line genome per se as discussed above.
Stress line did not significantly affect the times of HE or FE from the HWB. These results were also unexpected, because they did not confirm our previous studies (Satterlee and Jones, 1995; Jones et al., 1999) that found LS quail emerged into the unfamiliar and lighted compartment of the HWB apparatus sooner than did HS birds. Furthermore, only 7 to 9 d earlier in the TI testing of full siblings to the presently tested HWB quail, LS quail were found to be more resistant to induction into TI, and they had a shorter LATHEAD once successfully induced into TI than did HS quail. And, if the robust vocalization line differences that occurred in the dark compartment of the HWB truly reflect greater alarm calling in HS than LS quail as we have proposed, then it is logical to suspect HS quail to have been more frightened than LS quail and therefore for HS quail to take more time to show HE and FE. It is also widely held that fear is associated with inactivity in birds (Jones, 1987, 1996). In fact, in addition to the previous HWB emergence tests (Satterlee and Jones, 1995; Jones et al., 1999), when various locomotion behaviors were used as assessment elements in other behavioral tests of fear, HS quail have invariably (until now) shown reduced locomotor activity (e.g., various open field behaviors, Jones et al., 1992a; struggling in a crush cage, Jones and Satterlee, 1996, Jones et al., 2000).
Although we cannot offer a readily apparent explanation as to why no line differences in HE and FE behavior were presently detected, we did, however, find the times of both HE and FE from the HWB to be stymied, regardless of line, in the offspring of hens implanted with B. This is a new and important finding that lends support to the report by Janczak et al. (2006) that chicks hatched from eggs injected with B show greater fear of humans. These latter findings also argue back in support of a transovarian link between B and heightened fear as measured by offspring inactivity. On the other hand, considering that B-implant treatment was presently ineffective in altering TI behaviors, and in lieu of the controversial literature on maternal-in ovo-B influences on altering fear behavior of offspring, the dilemma remains that these relationships are not straightforward and may or may not be dependent on in ovo B intervention.
In summary, the quail stress line genome may or may not be affecting certain fear and alarm responses in chicks via the same mechanism(s) that underlie(s) how elevating maternal B increases egg levels of B that in turn alter the fear behavior of progeny. Additional research will be needed to help clarify these issues.
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ACKNOWLEDGMENTS |
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FOOTNOTES |
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Received for publication February 22, 2008. Accepted for publication March 28, 2008.
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REFERENCES |
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