Impact of social stress during gestation and environmental enrichment during lactation on the maternal behavior of sows

The impact of a social stress in gestation and an enriched pen in lactation on components of sow maternal behavior was studied in a 2 × 2 factorial experiment. At breeding, 41 sows were assigned to a social mixing stress treatment (T) during mid-gestation or a control group (C). During lactation, half of the T and C sows were housed in straw enriched pens (E) (1.57 m × 4.10 m) and the others in standard farrowing crates (S) (0.68 m × 2.10 m). The mixing stress consisted in introducing each T sow to the home pen of two unfamiliar sows twice for 1 week, from d 39 to 45 and 59 to 65 of gestation. Aggressive behavior was observed and lesion scores were taken to confirm that a social stress occurred. During lactation, the responses of sows to a simulated piglet crush test on d 3 and an isolated piglet playback test on d 21 were observed. Postural budgets of sows were automatically detected using accelerometers on d 5 and 19 of lactation. Sow-initiated social contacts with the piglets were observed continuously from video recordings on d 6 and 20 of lactation. Data were analyzed with a mixed models procedure. The social stress treatment had an impact on the response of sows to isolated piglet vocalizations with T sows showing longer latencies to respond vocally than C sows (P = 0.035). In early lactation, T sows spent more time lying ventrally than C sows (P = 0.007). Furthermore, the social stress had an impact on the space use in the enriched housing, with T sows spending less time in the nesting straw area of the pen than C sows (P = 0.018). Housing also impacted maternal behavior with E sows tending to spend more time lying ventrally than S sows in late lactation (P = 0.067) and tending to have more social contacts with their piglets than S sows in early lactation (P = 0.058). In conclusion, the social stress during gestation had a slight negative impact on sow maternal behavior, and while an enriched farrowing pen allowed for more opportunities to express maternal behavior, it did not counteract the negative effects of gestation stress.     

Chronic maternal stress affects growth, behaviour and hypothalamo-pituitary-adrenal function in juvenile offspring

Maternal stress during pregnancy, particularly that combined with low socioeconomic status (SES), has been linked to an increased risk for impaired behavioural and emotional development and affective disorders in children. In animal models, acute periods of prenatal stress have profound effects on hypothalamo-pituitary-adrenal (HPA) function and behaviour. However, few studies have determined the impact of chronic exposure to stress in animal models. The objective of this study was to determine the effects of chronic maternal stress (CMS) during the 2nd half of pregnancy and nursing on growth, locomotor behaviour and HPA axis function in juvenile guinea pig offspring. Pregnant guinea pigs were exposed to a random combination of variable stressors every other day over the 2nd half of gestation and from postnatal day (pnd) 1 until weaning (pnd25). CMS mothers displayed increased basal salivary cortisol levels in the later stages of pregnancy compared to control mothers (p < 0.05). The male offspring of CMS mothers had a lower bodyweight, which was maintained to weaning (p < 0.01). In open-field testing, CMS male offspring showed a decrease in activity compared to controls (p < 0.05). There was no effect of CMS on bodyweight or activity in female offspring. In contrast, both male and female offspring born to CMS mothers displayed increased (p < 0.05) basal salivary cortisol at pnd25, but a blunted adrenocortical response to exposure to the novel open-field enclosure. In conclusion, CMS leads to modification of growth trajectory, locomotor activity and adrenocortical responses to stress in juvenile offspring. Further, males appear considerably more vulnerable to these effects than females.     

Effects of environmental enrichment on reproduction

Although there have been few demonstrations of a direct empirical relationship between environmental enrichment and reproductive success in captive animals, indirect and anecdotal evidence indicates the importance of physical and temporal complexity for reproduction. We discuss three major mechanisms through which environmental enrichment that specifically increases the complexity of an animal's surroundings may influence reproductive physiology and behavior: developmental processes, modulation of stress and arousal, and modification of social interactions. In complex environments developing animals learn that performing active behavior produces appropriate functional outcomes. Learning to control their environment influences their ability to adapt to novel situations, which may profoundly influence their reproductive behavior as adults in breeding situations. Chronic stress may compromise reproductive physiology and behavior; enrichment reduces stress by providing increased opportunity for behavioral coping responses. However, some degree of acute stress may be beneficial for reproduction by maintaining an animal's level of responsiveness to socio-sexual stimuli necessary for sexual arousal and reproductive activation. Finally, environmental enrichment may influence reproductive success by stabilizing social groups, reducing aggression and increasing affiliative and play behaviors. It is concluded that multi-variate multi-institutional behavioral research in zoos will play an increasingly important role in the successful captive propagation of many species by closely examining relationships between environmental variables and reproductive potential of individual animals. © 1994 Wiley-Liss, Inc.     

Effects of BCH 325 (Pro-D-Phe-Pro-Gly) on open field behavior after chronic stress procedure.

The chronic stress model is an animal model relevant to clinical depression. The chronic stress rat group showed a significant decrease in motor activity. The reduced open field activity could be restored by subchronic treatment with imipramine and doxepine. Subchronic treatment with haloperidol, amphetamine and methylglucamine orotate was not able to alter the reduced open field activity. Previous treatment with a beta-casomorphin derivative (BCH 325) significantly antagonized the effect of chronic stress. The findings point out an antidepressant-like effect of BCH 325.     

Effects of pregnancy and chronic exercise on maternal cardiac structure and function

This study examined the interactive effects of pregnancy and aerobic conditioning on maternal cardiac structure and function. Effects of closely monitored cycle ergometer conditioning were studied during the second (TM2) and third trimesters (TM3) in 22 previously sedentary pregnant women (exercised group, EG) and a nonexercising pregnant control group with similar characteristics (CG, n = 19). Subjects were studied in the resting state by two-dimensional echocardiography and during cycle ergometer exercise at three steady-state power outputs at the start of TM2 (ENTRY), at the end of TM2 and TM3 (postconditioning), and 3-4 months postpartum (NPR, nonpregnant reference, CG only). Aerobic conditioning did not increase left ventricular dimensions beyond those attributable to pregnancy itself. In addition, in contrast with previous studies of nonpregnant women, physical conditioning during pregnancy did not reduce heart rate (HR) in the resting state. During exercise, the slope of the HR versus oxygen uptake (VO2) regression decreased significantly between preconditioning and the end of TM3 in the EG, suggesting that training-induced reductions in HR become more evident with increasing exercise intensity. Also, significant reductions in oxygen pulse (VO2/HR) were observed at all three work rates in the CG, but not in the EG. These findings support the hypothesis that the cardiovascular effects of aerobic conditioning are obscured by more powerful effects of pregnancy in the resting state but become "unmasked" during strenuous exercise.     

The influence of environmental enrichment on Chinese visitor behavior

Welfare improvements for nonhuman animals should aim to satisfy the needs of visitors as well as those of the animals. Little research has been conducted, however, and existing work is confined to zoos in developed countries. This article reports the behavioral responses of Chinese visitors to environmental enrichment improvements in a zoo enclosure. Visit, viewing, and stopping behaviors significantly increased at the transformed exhibit, indicating that it provoked greater visitor interest. Furthermore, increased intragroup behaviors suggested that the exhibit probably motivated visitors to interact socially. The positive impact of the exhibit changes supports the enrichment efforts taking place in zoos around the world. The changes also provide encouragement for zoos in developing countries such as China because greater visitor interest provides a strong argument and an incentive for improving welfare standards.     

The effect of housing and environmental enrichment on stereotyped behavior of adult vervet monkeys (Chlorocebus aethiops)

Little information is available on the response of vervet monkeys to different housing conditions or on the suitability of enrichment devices or methods for vervet monkeys. In this study, the authors evaluated the occurrence of stereotyped behavior in adult vervet monkeys under various conditions of housing and enrichment. The variables included cage size, cage level (upper or lower), enrichment with a foraging log, enrichment with an exercise cage and presence of a mate. The authors first determined the incidence of stereotyped behavior in captive-bred, singly housed adult female and male vervet monkeys. They then exposed monkeys to different housing and enrichment situations and compared the incidence of stereotyped behavior among the monkeys. The authors found that more females than males engaged in stereotyped behavior and that females, on average, engaged in such behavior for longer periods of time than males. Stereotyped behavior was most often associated with a small, single cage. The average amount of observed stereotyped activity in monkeys housed in a small cage was significantly lower when the monkeys had access to either a foraging log or an exercise cage. Stereotyped behavior was also lower in female monkeys that were housed (either with a male or without a male) in a larger cage. The least amount of abnormal behavior was associated with the largest, most complex and enriched housing situation. Males and females housed in cages on the lower level of two-level housing engaged in more stereotyped behavior than did monkeys housed in the upper level, regardless of the presence or type of enrichment provided.     

Transient sex-related changes in the mice hypothalamo-pituitary-adrenal (HPA) axis during the acute phase of the inflammatory process

The potential role of endogenous sex hormones in regulating hypothalamo-pituitary-adrenal (HPA) axis function was investigated after a single injection of endotoxin in adult (8 week old) BALB/c mice of both sexes. The effect of LPS on plasma ACTH, corticosterone (B), testosterone and oestradiol (E) levels and on anterior pituitary (AP) ACTH and adrenal B contents at different times after treatment was studied. The results indicate that: (a) basal B but not ACTH plasma levels were significantly higher in female than in male mice; (b) LPS significantly increased both ACTH and B plasma levels over the baseline 2 h after injection, both hormone levels being higher in female than in male mice; (c) although plasma ACTH concentrations recovered the basal value at 72 h after LPS in animals of both sexes, plasma B levels returned to the baseline only at 120 h after treatment; (d) E plasma levels significantly increased 2 h after LPS and returned to the baseline at 72 h post-treatment, in both sexes; (e) at 2 h after LPS, testosterone plasma levels significantly decreased in male mice and increased in female mice, recovering the baseline level at 120 and 72 h after LPS, respectively; (f) AP ACTH content was similar in both sexes in basal condition and it was significantly diminished 72 h post-treatment without sex difference; whereas AP ACTH returned to basal content 120 h after LPS in males, it remained significantly decreased in females; (g) basal adrenal B content was higher in female than in male mice, and it significantly increased in both sexes 2 h post-LPS, maintaining this sex difference. Whereas adrenal B returned to basal content 72 h after treatment in male mice, it remained significantly enhanced up to 120 h post-LPS in female animals. The data demonstrate the existence of a clear sexual dimorphism in basal condition and during the acute phase response as well as in the recovery of the HPA axis function shortly after infection.     

Effects of selected behavioral enrichment devices on behavior of Western Lowland Gorillas (Gorilla gorilla gorilla)

Environmental complexity plays an integral role in the activity and psychological well-being of primates. The experiment described in this article evaluates the effects of nonintrusive, inexpensive, and easily managed behavioral enrichment devices on the behavior of a group of captive Western lowland gorillas. Devices used included cardboard boxes containing food items, paper bags containing food items, burlap rags, and willow and maple browse. The enrichment devices increased foraging, social play, and solitary play behaviors. Sedentary behaviors decreased. Rags, bags, browse, and boxes did not statistically decrease the incidence of regurgitation/reingestation or coprophagy. Depending on the type of enrichment item used, the effects on agonism and manipulation of enrichment item were variable. To make informed management decisions about the psychological well-being of captive animals, it is important to objectively quantify and examine the influences on their behavior.     

A hypothetical index of hypothalamic-pituitary-adrenal (HPA) axis function.

The short ACTH stimulation test (SST) is commonly used by clinicians to determine the integrity of the hypothalamic-pituitary-adrenal (HPA) axis in cases of suspected adrenal insufficiency. Concern has arisen for under-diagnosis of central hypocortisolism using the standard dose (250 mcg) SST with the recent introduction of low-dose (1 mcg) SST as a potentially more sensitive test for central hypocortisolism. Amidst this uncertainty, I will present a novel index hypothesized to extend the sensitivity of the standard dose SST as an adjunctive diagnostic tool for any level of hypocortisolism.     

Effects of environmental features on flocking behavior

Social animals or insects in nature often exhibit a form of emergent collective behavior known as flocking. Animal flocks are strongly affected by the spatial heterogeneity representing static environmental factors, such as landscape. To investigate how this heterogeneity affects flocking behavior, I constructed a two-dimensional, lattice-based environment, in which the spatial heterogeneity was generated using a neutral landscape model. A value ranging from 0.0 to 1.0 was assigned to each lattice site. High values represent favorable environmental conditions, and thus, individuals tend to be attracted to the area with the highest values. The value H controlled the degree of heterogeneity and the value h represented the amplitude of the heterogeneity. To characterize the effect of the spatial heterogeneity, I introduced the order parameter, ?, defined as the absolute value of the average unit velocity. When all individuals move in the same direction, the value of ? is 1, while the value of ? becomes 0 in the case where all individuals move randomly. In this simulation, the flock size, N, has the values of 20, 40, 60, 80 and 100. I investigated the average value of ? and ?_s, for 1500 < t < 2000. The simulation results showed that the value of ?_s increased with an increase in H, regardless of N, until becoming saturated. When the value of h was higher, the value of ?_s became lower.     

Effects of maternal stress coping style on offspring characteristics in rainbow trout (Oncorhynchus mykiss)

Maternal size, age, and allostatic load influence offspring size, development, and survival. Some of these effects have been attributed to the release of glucocorticoids, and individual variation in these stress hormones is related to a number of traits. Correlated traits are often clustered and used to define the proactive and reactive stress coping styles. Although stress coping styles have been identified in a number of animal groups, little is known about the coupling between stress coping style and offspring characteristics. In the present study, plasma cortisol levels in ovulated mothers and cortisol levels in non-fertilized eggs from two rainbow trout (Oncorhynchus mykiss) strains selected for high (HR) and low (LR) post-stress plasma cortisol levels were compared. Offspring characteristics such as egg size, larval growth, and energy reserves also were compared between the two strains. Maternal plasma and egg cortisol levels were correlated, but no difference between the HR and LR strains was detected in either parameter. LR females produced larger eggs, and larvae with larger yolk sacs compared to HR females, however no differences in larval body size (excluding the yolk) was detected between strains. Considering that the HR and LR strains have a number of correlated behavioral and physiological traits that resemble the reactive and proactive stress coping styles, respectively, the results suggest that proactive mothers invest more energy into their offspring, producing larvae with larger energy reserves. It is possible that larger energy reserves in proactive larvae support the energy requirement for establishing and defending territory in salmonid fish. Furthermore, in the present study we found a positive relationship between mother plasma cortisol and egg cortisol; however neither mother plasma cortisol nor egg cortisol differed between strains. These results indicate that cortisol endowment from the mother to the offspring plays a minor role in the transfer of the behavioral and physiological traits which separates these strains.     

Effects of environmental stress or ACTH treatment during pregnancy on maternal and fetal plasma androstenedione in the rat

Prenatal stress applied during the last trimester of pregnancy has been shown to alter fetal development and influence adult sexual behavior. Since androstenedione (Δ⁴) has the potential to participate in differentiation processes, this study was designed to assess the effect of prenatal stress on maternal and fetal Δ⁴ titers. Restraint/illumination/heat (environmental stress) or ACTH injections were used to stress pregnant rat dams beginning on Day 14 of pregnancy. Blood samples and organ weights were obtained from nonpregnant animals, pregnant rats on Days 5, 10, 15, 18, and 20 of pregnancy, and fetuses on Days 18 and 20 of gestation. Maternal and male and female fetal Δ⁴ titers were determined by radioimmunoassay. ACTH and environmental stress significantly reduced fetal body weight and male anogenital distance. Environmental stress also significantly reduced the size of 20-day fetal adrenals and testes. Each treatment caused significant short-term (1 hr after treatment) and long-term (16 hr after treatment) elevation of maternal plasma Δ⁴ on Days 15 and 18 of gestation, but only short-term elevation of Δ⁴ titers on Day 20. ACTH treatment did not cause long-term elevation of fetal Δ⁴ although both ACTH treatment and environmental stress generated a significant short-term increase in fetal Δ⁴ titers. Environmental stress produced long-term elevation of fetal Δ⁴ in 18-day fetuses of both sexes and in 20-day female fetuses. It is concluded that maternal stress and exogenous ACTH significantly elevate maternal and fetal Δ⁴ titers during the prenatal period postulated to be critical in sexual differentiation of the rat brain.     

Bobcat (Lynx rufus) breeding in captivity: the importance of environmental enrichment

Environmental enrichment is an improvement in the biological functioning of nonhuman animals in captivity resulting from modifications to their environment; however, specifying appropriate and practical measures of enrichment is problematic. This study analyzes the behavior of 4 bobcats (Lynx rufus) in the Jerez Zoo before and after the application of a global program of environmental enrichment that included (a) changes in the size and complexity of their installations, (b) the introduction of new objects into compounds, (c) changes in diet, and (d) modifications in the grouping of animals. A factorial correspondence analysis showed a highly significant relationship among individual animals, behavior, and experimental design. Behaviors such as locomotion, repeated pacing, vigilance, and grooming more often occurred before enrichment, whereas exploratory and food behaviors were more often associated with the enrichment phase. After the implementation of the enrichment program, the bobcats bred successfully for the first time since their arrival in the zoo.     

Maximizing the effectiveness of environmental enrichment: Suggestions from the experimental analysis of behavior

Environmental enrichment programs provide benefits to both captive animals and the facilities that house them, but cost time and resources to design, implement, and maintain. As yet, there have been few theoretically based guidelines to assist animal care staff in establishing cost-efficient enrichment methods that both elicit the desired behavioral changes and maintain their success over time. We describe several well-studied principles from the field of experimental analysis of behavior, including intrinsic reinforcement, extrinsic reinforcement, habituation, extinction, and schedules of reinforcement that could be very useful for evaluating the short- and long-term effectiveness of enrichment. We use this theoretical framework to generate testable hypotheses and provide examples of enrichment studies relevant to our predictions. In particular, we suggest that enrichment devices that offer extrinsic reinforcement (food, social access, etc. as a result of performing behaviors) should produce greater and more prolonged changes in behavior than devices that rely on the behavior itself being reinforcing to the animal. For techniques that provide no extrinsic reinforcement, using stimuli that are novel, are more different from the environment, have been withheld or altered in some way, or are presented less frequently may help reduce habituation. For techniques that provide extrinsic reinforcement, making reinforcement more difficult to obtain and providing more or higher quality reinforcers may increase the long-term success of the enrichment program. In addition, enrichment may be more effective if animal care staff avoid continuously reinforcing behaviors after they are established, enriching immediately after feeding, and exposing animals to enrichment when reinforcement is no longer available. While the current enrichment literature supports the application of behavior analytic theory, empirical evaluation of many of our predictions is still needed.     

Effects of nutritional enrichment on the production of acetone-butanol-ethanol (ABE) by Clostridium acetobutylicum

Clostridium acetobutylicum is an industrially important organism that produces acetone-butanol-ethanol (ABE). The main objective of this study was to characterize the effects of increased cell density on the production of ABE during the phase transition from acidogenesis to solventogenesis in C. acetobutylicum. The increased ABE productivity of C. acetobutylicum was obtained by increasing the cell density using a newly designed medium (designated C. a cetobutylicum medium 1; CAM1). The maximum OD₆₀₀ value of C. acetobutylicum ATCC 824 strain obtained with CAM1 was 19.7, which is 1.8 times higher than that obtained with clostridial growth medium (CGM). The overall ABE productivity obtained in the CAM1-fermetation of the ATCC 824 strain was 0.83 g/L/h, which is 1.5 times higher than that (0.55 g/L/h) obtained with CGM. However, the increased productivity obtained with CAM1 did not result in an increase in the final ABE titer, because phase transition occurred at a high titer of acids.     

Influence of environmental enrichment and handling on the acute stress response in individually housed mice

In this study we investigated the effect of environmental enrichment and handling on the acute physiological stress response caused by short periods of restraint in individually housed female mice. Heart rate (HR) and body temperature (BT) were measured by radiotelemetry and compared with plasma corticosterone (pCORT) levels. Also, postmortem thymus weight and tyrosine hydroxylase (TH) activity were assessed. The acute stress response was seen in both HR and BT. Enrichment and handling were found to increase rather than decrease this stress response, but pCORT values, measured 90 min after restraint, suggested a lower stress response in the enriched groups. No effect was found with thymus weight or TH as parameters.