Regulation of dustbathing in feathered and featherless domestic chicks: the Lorenzian model revisited

In Lorenz's 'psychohydraulic model', behaviour is regulated by performance: the motivation to perform a behaviour builds up with time and can be reduced only by performance itself. However, a convincing example of Lorenzian regulation has been lacking. We studied dustbathing in featherless and feathered chicks, Gallus gallus domesticus, the latter trained to dustbathe on glass with sand underneath. In both cases the function, that is, plumage cleaning, was eliminated. In both featherless and feathered chicks dustbathing increased with time since the last performance as found in normal dustbathing, and there was a compensatory adjustment when the amount of previous dustbathing was artificially reduced. We suggest that the amount of dustbathing performed at any one time may be controlled by the cooperative action of a deprivation or timer mechanism and a counter mechanism. The timer mechanism responds to the time since the last performance, whereas the counter mechanism records the number of elements during that last dustbathing. Alternatively, there may be a single mechanism that continually records the deficit in the amount of dustbathing performed. Lorenz's model may be valid not only for dustbathing in fowl but also for other comfort behaviours. Our results suggest the existence of behavioural needs that can be satisfied only by performance. Copyright 1999 The Association for the Study of Animal Behaviour.     

Responses to pulsed light stimuli in a pulmonate slug (Limax pseudoflavus)

The phase changes in the activity rhythm of the pulmonate slug Limax pseudoflavus caused by light pulses (30 min, 10,000 lux) in otherwise dark conditions were recorded using time lapse cinematography and tipping aktographs. From the timing of the pulse relative to an individual activity cycle a phase response curve was constructed. Phase delays occurred when the pulse was administered in the early subjective night and subjective day, phase advances were recorded during the mid and late subjective night.
The effects of two pulses (30 min, 10,000 lux) forming symmetric and asymmetric skeleton photoperiods were also recorded and related to the phase response curve. Both stable and unstable entrained states were found, the condition being dependent upon the relative timing of the two pulses and the previous activity onset. It was also shown that there was a time lag between the light pulse and its expressed phase setting effect. Thus phase setting is not instantaneous as with some insects.     

Circadian Changes in Anxiolysis-Related Behavior of Syrian Hamsters. Correlation with Hypothalamic GABA Release

The diurnal variations in anxiolysis and exploratory behavior were examined in a plus-maze paradigm in Syrian hamsters exposed to 14 h light: 10 h dark photoperiods or to constant darkness for 3 days. The percent of time spent in open arms and the percent of entries to open arms (both indexes of anxiety-related behavior) as well as the number of crosses to both arms (an index of locomotor behavior) showed significant daily variations under the two lighting conditions, maxima being found at night (2400-0400 h). Flumazenil (5 mg/kg) injected at the middle of the light (at 1600 h) or dark period (at 0400 h) decreased by 39-54 % anxiolysis-related behavior without affecting locomotor activity significantly. 3 H-?-aminobutyric acid (GABA) release from preoptic-medial basal hypothalamic explants obtained from hamsters exposed to 14 h light: 10 h dark photoperiods attained maximal values at 2400-0400 h. The results further supported the existence of circadian changes in anxiolysis-related behavior in Syrian hamsters that correlated with an increased hypothalamic GABA release.     

Photosensitivity in body mass and testicular activity of brahminy myna, Sturnus pagodarum

This study analyzed photoperiodic sensitivity of brahminy myna (Sturnus pagodarum), which is a seasonally breeding bird species. During regression phase of the reproductive cycle (in early September), groups of myna were exposed to artificial photoperiods that were either close to or longer than those brahminy myna experiences at this time in wild at 29 degrees N. Following a 14-week exposure to such photoperiods (hours of light: hours of darkness; 13L:11D, 12L:12D, 11.5L:12.5D and 11L:13D), the birds were subjected to a longer day length (16L:8D) for another 9 weeks to test whether pre-treatment with varying photoperiods had an effect on subsequent long day photostimulation. There was a progressive increase in body mass under different pre-treatment photoperiods, with a faster increase in 11L:13D and 11.5L:12.5D than in 12L:12D and 13L:11D. When subjected to 16L: 8D, however, all groups showed decline in body mass. By contrast, the testes were not stimulated under 11L:13D and 11.5L:12.5D, and only one individual of the groups under 12L:12D and 13L: 11D showed a small testis recrudescence. All birds except one individual of the 13L:11D group, however, showed testis recrudescence when subjected to 16L:8D. These results suggest that body mass and testes have a different profile of photoperiodic response and appear to have two different threshold photoperiods for dissipating the post-reproductive refractoriness.     

Effects of light (photoperiod,spectral composition) on the population dynamics of Tisbe holothuriae Humes (Copepods,Harpacticoida)

The harpacticoid copepod Tisbe holothuriae Humes was reared for several generations under different photoperiods (white light) and spectral quality (continuous illumination). The main life cycle parameters were measured and demographic variables were determined. Development time was retarded under green wavelengths and at photoperiods different from LD 12:12, especially under continuous white light. Total body length of females and males was shortest under LD 12:12 and longest under constant dark. Photoperiods different from LD 12:12 prolonged the interval between broods of egg sacs. Short photoperiods caused higher abortion rate, while long photoperiods retarded the maturation time of the egg sacs. Red wavelengths stimulated the abortion rate. Photoperiod LD 12:12 and blue wavelengths increased the production of offspring and the life span of females. Highest intrinsic rates of natural increase were estimated under LD 12:12 (rm =0.304) and blue wavelengths (rm = 0.254). These light conditions proved the most favourable for rearing Tisbe holothuriae.     

Effect of photoperiod on incubation period in a wild passerine,Sylvia atricapilla

Time required for avian embryos to develop is influenced by incubation temperature and the amount of time adults incubate eggs. Experiments on poultry indicate that photoacceleration, the light‐induced stimulation of embryonic development, decreases the length of the incubation period as embryos receive more light. We hypothesized that eggs of wild birds exposed to longer periods of light should also have shorter incubation periods. We tested whether photoacceleration would occur in a species of open‐cup nesting passerine, the blackcap Sylvia atricapilla. We artificially incubated blackcap eggs under four different photoperiods, four hours of light (4L) and 20 h of dark (20D), 12L:12D, 20L:4D, and a skeleton photoperiod (1 h light, 2 times per day) that framed a 20 h day. While incubation periods were accelerated with increasing photoperiod length, the differences among photoperiods of 4, 12 and 20L were weak. Embryos exposed to skeleton photoperiods developed as fast as those exposed to 20L and significantly faster than those exposed to 4L and 12L treatments. Skeleton photoperiods may most closely approximate natural patterns of light exposure that embryos experience during dawn and dusk incubation recesses typically associated with adult foraging. If our results from this species also occur in other wild birds, exposure to different day lengths may help explain some of the variation in the observed seasonal and latitudinal trends in avian incubation period.     

The effect of various photoperiods on daily oscillations of serum corticosterone and insulin in rats.

The effect of various photoperiods on circadian rhythms of chosen parameters was investigated in laboratory rats. SPF male Wistar rats were adapted for six weeks to artificial light-dark cycles (LD 8:16, 12:12, 16:8). The light was switched on at 07.00 h in all regimens. The rats were killed at 3-hour intervals within 24 h, the serum concentration of corticosterone, insulin, glucose, food and water intake was determined. The external and computative acrophases of corticosterone varied in every photoperiod being dependent on the duration of light, the mesor values decreased in LD 16:8 in comparison with other photoperiods. The external acrophase of insulin was located 4 h after light onset in LD 8:16 and 12:12, in LD 16:8 one hour before light onset. The mesor values were approximately equal in all photoperiods. The circadian rhythms of glucose were similar in all regimens. Circadian variation of food and water consumption culminated at the same time in all regimens, the amount of food consumed in light increased with the light duration. Various photoperiods remarkably influenced circadian oscillations of corticosterone and in part food and water intake which could be considered as photoperiodic traits.     

Photoinhibition of photosystems I and II using chlorophyll fluorescence measurements

In this study the photoinhibition of photosystems (PS) I and II caused by exposure to high intensity light in oat (Avena sativa, var Prevision) is measured by the emission of chlorophyll fluorescence in intact leaves adapted to darkness. The maximal quantum yield of PS II was lower in plants grown under high light intensity than in plants grown under low intensity, which indicates that PS II is photoinhibited by such conditions. PS I was more stable than PS II in plants exposed to strong light for a moderate time (five photoperiods) since the oxidised plastoquinone pool size under far-red (FR) light was similar in plants grown under high light intensity to plants grown under low intensity, probably as a result of the cyclic electron flow around PS I being stimulated in response to high light intensity. However, over longer times (10 photoperiods) the PS I was photoinhibited, since the oxidised plastoquinone pool size under FR light increased as a consequence of the decrease in PS I activity caused by high light intensity. This practical is intended for advanced students of plant biochemistry and plant physiology.     

Food-Anticipatory Activity in Syrian Hamsters: Behavioral and Molecular Responses in the Hypothalamus According to Photoperiodic Conditions

When food availability is restricted, animals adjust their behavior according to the timing of food access. Most rodents, such as rats and mice, and a wide number of other animals express before timed food access a bout of activity, defined as food-anticipatory activity (FAA). One notable exception amongst rodents is the Syrian hamster, a photoperiodic species that is not prone to express FAA. The present study was designed to understand the reasons for the low FAA in that species. First, we used both wheel-running activity and general cage activity to assess locomotor behavior. Second, the possible effects of photoperiod was tested by challenging hamsters with restricted feeding under long (LP) or short (SP) photoperiods. Third, because daytime light may inhibit voluntary activity, hamsters were also exposed to successive steps of full and skeleton photoperiods (two 1-h light pulses simulating dawn and dusk). When hamsters were exposed to skeleton photoperiods, not full photoperiod, they expressed FAA in the wheel independently of daylength, indicating that FAA in the wheel is masked by daytime light under full photoperiods. During FAA under skeleton photoperiods, c-Fos expression was increased in the arcuate nuclei independently of the photoperiod, but differentially increased in the ventromedial and dorsomedial hypothalamic nuclei according to the photoperiod. FAA in general activity was hardly modulated by daytime light, but was reduced under SP. Together, these findings show that food-restricted Syrian hamsters are not prone to display FAA under common laboratory conditions, because of the presence of light during daytime that suppresses FAA expression in the wheel.     

Induction of flowering in the duckweed Lemna paucicostata under non-inductive photoperiods by tannic acid

Flowering in Lemna paucicostata 6746 could be induced by tannic acid under strictly non-inductive photoperiods. This polyphenol completely abolished the photoperiodic sensitivity of strain 6746 as flowering could also be obtained under continuous light (nearly 80% flowering was recorded in the plants supplied with 10⁻⁵ tannic acid). Though its mode of action is unknown, tannic acid is unlikely to act as a gibberellin-antagonist in its effect on flowering in strain 6746.     

Room novelty, sex, scopolamine and their interactions as determinants of general activity and rearing, and light-dark preferences in rats

Male and female rats were assessed for effects of scopolamine on general activity, rearing and light-dark preferences when tested in either a familiar or a novel room. Males but not females reared more often when tested in the familiar rather than novel room, and the response was increased by scopolamine for all rats combined. Whereas scopolamine increased general activity for females (but not males) in the familiar room, it decreased the response for males (but not females) in the novel room. Females crossed more often between the dark and light sides of a light-dark box and, when treated with saline but not drug, spent more time in the light side than males. Scopolamine reduced the amount of time spent in the light side for females only. While the results were discussed mainly in terms of sex differences in fearfulness, their principal value was in demonstrating the effectiveness of room novelty and sex in determining levels of the behaviors recorded, and drug responsiveness.     

Dustbathing as a regulatory mechanism

Three mathematical models were developed to describe how some species of birds, such as bobwhite quail (Colinus virginianus) regulate the amount of oil on their plumage. The models assume that dustbathing plays a significant role in this regulatory process. They differed primarily in their assumptions about the relationship between oiling and dustbathing behavior. Several experiments were run to check on the implications of the models.     

An evaluation of potential dustbathing substrates for commercial broiler chickens

Provision of an appropriate dustbathing substrate may allow broiler chickens to satisfy a natural motivation and give them an opportunity to exercise. The main aim of this study was to evaluate the extent to which different substrates promote dustbathing behaviour in broilers. The trial was replicated over three production cycles in one commercial broiler house, with ~22 000 Ross broilers housed per cycle. The birds were provided with access to five experimental substrates from day 10 of the 6-week production cycle. The substrates included the following: (1) peat (P), (2) oat hulls (OH), (3) straw pellets (SP), (4) clean wood shavings (WS), and (5) litter control (C). The substrates were provided in 15 steel rings (1.1 m in diameter, three rings per substrate) dispersed throughout the house. The level of occupancy of the rings, behaviours performed in each substrate, and the effect of ring position (central or edge of house) were assessed in weeks 3, 4, 5 and 6 using scan sampling from video footage. Where substrates successfully promoted dustbathing, the length and components of the bouts (including number of vertical wing shakes and ground pecks) were also assessed. Results showed that birds used P significantly more than the remaining substrates for dustbathing (P<0.001). Oat hulls were the second most preferred substrate for dustbathing, with significantly more birds dustbathing in the OH compared with SP, WS and C (P<0.001). The least sitting inactive was also seen in the P and OH rings compared with the SP, WS and C (P<0.001). The highest levels of foraging were recorded in the P, OH and WS compared with SP and the C. Position of the rings did not affect the types of behaviours performed in any substrate, although overall more birds were counted in the central compared with edge rings (P=0.001). More detailed information on dustbathing behaviour was only recorded in the P and OH treatments, and there were no differences in the length of dustbathing bout, or components of the bout between them (P>0.05). The use of OH is likely to be more environmentally sustainable than that of P, and our results suggest that this substrate is relatively successful in promoting dustbathing. However, a preference was still observed for P and further work should investigate whether other suitable substrates could better reflect its qualities.     

Effect of photoperiod manipulation on the growth performance of juvenile lenok,Brachymystax lenok (Pallas, 1773)

The effect of four different light regimes on growth was studied in lenok, Brachymystax lenok. Fish with average weights of 5.5 g were subjected to four different photoperiods (0L:24D, 6L:18D, 12L:12D and 24L:0D) for 35 days. The specific growth rate (SGR) of lenok in 24‐h darkness had a significantly higher SGR than those in the continuous light regime (P < 0.05); however, there was no significant difference among fish exposed to 6L:18D, 12L:12D and 24L:0D photoperiods. There was a tendency for higher food intake over the light period extension from 0L to 24L, and feed intake was significantly higher in the continuous light group than in 24‐h darkness (P < 0.05). No significant difference in feed conversion efficiency (FCE) was observed between fish exposed to 0L:24D and 6L:18D photoperiods, however, the FCE in both photoperiods was significantly higher than that in the other two groups. The final survival rate of juveniles varied from 79.67 to 95.33%, with significant differences among experimental groups. Fish tested in continuous illumination spent much more energy on respiration and excretion while depositing less energy for growth than in the other photoperiods. In contrast, fish in 24‐h darkness deposited more energy for growth and spent less energy on respiration and excretion. Results show that photoperiod manipulation can affect growth, and that a continuous dark regime could improve growth in lenok at this stage of development.     

Influence of Photoperiod, Daylength, and Feeding Schedule on Tadpole Growth and Development

The metamorphic rate of Rana pipiens tadpoles was studied under different photoperiods, daylengths, and feeding schedules. Tail resorption and hindlimb growth and development induced by immersion in 30 μg/l thyroxine (T₄) were accelerated under longer photoperiods and continuous light when 6L: 18D, 12L: 12D, 18L: 6D, and 24L regimes were compared. Constant light exposure did not produce faster development than an 18 hr photoperiod, and initially was less effective. The rate of spontaneous and T₄-induced metamorphosis was greater the shorter the day on 9L:9D, 12L: 12D, or 15L: 15D cycles, although all groups received the same overall amount of light, but in different dosages. When feeding schedule but not the LD cycle was varied, groups of tadpoles fed on 18, 24, or 30 hr regimes showed no differences in growth and development rate on 19L: 5D, and only random variations under continuous light. Differences in metamorphic rate on 18, 24, or 30 hr days are not due to the feeding schedules, but to the LD cycles. From these experiments we conclude that illumination, particularly the length and frequency of the photoperiod, affects the utilization of T₄. Development rates independent of the total amount of illumination, but related to daylength and light schedule, suggest interaction of light with an endogenous timing mechanism.     

Prenatal and postnatal effects of corticosterone on behavior in juveniles of the common lizard, Lacerta vivipara

Many animals exhibit dramatic responses when subjected to a stressor. A classic marker of the stress response is an increase in plasma glucocorticoids, but this constitutes only one step in the cascade from experience of a stressor to wider organismal changes, including behavior. The behavioral sensitivity to glucocorticoids would determine the consequences of the stress-related alteration of behavior for the organism. In this study we explored, under laboratory conditions, the prenatal and postnatal effects of corticosterone on activity and thermoregulation of juveniles of the common lizard, Lacerta vivipara. Activity was measured as the time spent moving and the time spent scratching the wall in an empty terrarium. Thermoregulatory behavior was measured as the time spent motionless under a light bulb. Activity and thermoregulation of juveniles of the common lizard showed a different sensitivity to prenatal and postnatal corticosterone treatment, modulated by juvenile sex and maternal condition. Prenatal corticosterone manipulation influenced the time spent moving in both sexes. By contrast, only juvenile females increased the time spent scratching the walls of the terrarium when corticosterone was delivered both at the prenatal and postnatal stage. Prenatal hormone manipulation increased the time spent basking by juveniles issued from large females. These results suggest that, in addition to influencing a variety of behavioral and morphological traits, corticosterone may also play an important role in the regulation of activity and thermoregulation of juvenile lizards, modulated by individual sex and maternal condition.     

Maternal behaviour and early behavioural ontogeny of grey seals (Halichoerus grypus) on the Isle of May, UK

Female grey seals allocated more resources to individual sons than to individual daughters. Mothers of male pups spent significantly larger proportions of their time defending and feeding their young than did mothers of female pups. The amount of time females spent ashore and the proportion of time spent in various behavioural states was influenced by the habitat of the birth site and the age of the pup.
Grey seal neonates spent the majority of their time prone and immobile, perhaps to allow maximum energy retention (maximum mass gain) and to facilitate mothers relocating their pups. The proportion of time pups spent in the various behavioural states was influenced by the whelping habitat. Pups became increasingly active with age. Males were significantly more aggressive than females and, at one site they performed nosing behaviour with their mothers significantly more often than did female neonates.     

Control of diapause induction by a change in photoperiod in Melanoplus sanguinipes

The effect of photoperiod on diapause induction, developmental time and body size was examined in Melanoplus sanguinipes, the lesser migratory grasshopper. Contrary to what is found in most insects, facultative diapause-egg production is found to be controlled by changing rather than constant photoperiods. In addition, developmental time is shown to be faster under short-day photoperiods than long-day photoperiods. And finally, body size is larger under long-day photoperiods and smaller under short-day photoperiods. Implications of these results for the regulation of the seasonal life cycle of M. sanguinipes in the field are discussed.     

Glutamic Acid Dehydrogenase Activity in Linseed as Influenced by Plant Age and Photoperiod

Glutamic acid dehydrogenase (GDH) activity was measured in linseed Linum usitatissimum ev. SH-1 at different growth intervals under normal and long photoperiodic conditions. In ungerminated seeds a low activity of GDH was recorded which increased rapidly during germination and thereafter decreased to a level when no GDH activity could be detected at 47 days of growth. An increase in its activity was recorded at 67, 87 and 107 days. Longer photoperiod (continuous light) increased the activity in roots markedly as compared to shorter photoperiods (normal days).     

Layer pullet preferences for light colors of light-emitting diodes

Light colors may affect poultry behaviors, well-being and performance. However, preferences of layer pullets for light colors are not fully understood. This study was conducted to investigate the pullet preferences for four light-emitting diode colors, including white, red, green and blue, in a lighting preference test system. The system contained four identical compartments each provided with a respective light color. The pullets were able to move freely between the adjacent compartments. A total of three groups of 20 Chinese domestic Jingfen layer pullets (54 to 82 days of age) were used for the test. Pullet behaviors were continuously recorded and summarized for each light color/compartment into daily time spent (DTS), daily percentage of time spent (DPTS), daily times of visit (DTV), duration per visit, daily feed intake (DFI), daily feeding time (DFT), feeding rate (FR), distribution of pullet occupancy and hourly time spent. The results showed that the DTS (h/pullet·per day) were 3.9±0.4 under white, 1.4±0.3 under red, 2.2±0.3 under green and 4.5±0.4 under blue light, respectively. The DTS corresponded to 11.7% to 37.6% DPTS in 12-h lighting periods. The DTV (times/pullet·per day) were 84±5 under white, 48±10 under red, 88±10 under green and 94±8 under blue light. Each visit lasted 1.5 to 3.2 min. The DFI (g/pullet·per day) were 27.6±1.7 under white, 7.1±1.6 under red, 15.1±1.1 under green and 23.1±2.0 under blue light. The DFT was 0.18 to 0.65 h/pullet·per day and the FR was 0.57 to 0.75 g/min. For most of the time during the lighting periods, six to 10 birds stayed under white, and one to five birds stayed under red, green and blue light. Pullets preferred to stay under blue light when the light was on and under white light 4 h before the light off. Overall, pullets preferred blue light the most and red light the least. These findings substantiate the preferences of layer pullets for light colors, providing insights for use in the management of light-emitting diode colors to meet pullet needs.