Body Fattening Research Articles

Seasonality in tropical birds.

The survival of offspring depends on environmental conditions. Many organisms have evolved with seasonality, characterized as initiation-termination-reinitiation of several physiological processes (i.e., body fattening, molt, plumage coloration, reproduction, etc). It is an adaptation for the survival of many species. Predominantly seasonal breeders use photoperiod as the most reliable environmental cue to adapt to seasonal changes but supplementary factors like temperature and food are synergistically involved in seasonal processes. Studies from diverse vertebrate systems have contributed to understanding the mechanism involved in seasonal reproduction at the molecular and endocrine levels. Long-day induced thyrotropin (thyroid-stimulating hormone) released from the pars tuberalis of the pituitary gland triggers local thyroid hormone activation within the mediobasal hypothalamus. This locally produced thyroid hormone, T3, regulates seasonal gonadotropin-releasing hormone secretion. Most of the bird species studied are seasonal in reproduction and linked behavior and, therefore, need to adjust reproductive decisions to environmental fluctuations. Reproductive strategies of the temperate zone breeders are well-documented, but less is known about tropical birds' reproduction and factors stimulating the annual breeding strategies. Here, we address seasonality in tropical birds with reference to seasonal reproduction and the various environmental factors influencing seasonal breeding.

Control and adaptability of seasonal changes in behavior and physiology of latitudinal avian migrants: Insights from laboratory studies in Palearctic-Indian migratory buntings.

Twice-a-year migrations, one in autumn and the other in spring, occur within a discrete time window with striking alterations in the behavior and physiology, as regulated by the interaction of endogenous rhythms with prevailing photoperiod. These seasonal voyages are not isolated events; rather, they are part of an overall annual itinerary and remain closely coupled to the other annual subcycles, called seasonal life history states (LHSs). The success of migration depends on appropriate timing of the initiation and termination of each LHS, for example, reproduction, molt, summer nonmigratory, preautumn migratory (fattening and weight gain), autumn migratory, winter nonmigratory (wnM), prevernal (spring) migratory (fattening and weight gain), and spring migratory LHSs. Migration-linked photoperiod-induced changes include the body fattening and weight gain, nocturnal Zugunruhe (migratory restlessness), elevated triglycerides and free fatty acids, triiodothyronine and corticosterone levels. Hypothalamic expression of the thyroid hormone-responsive dio2 and dio3, light-responsive per2, cry1,and adcyap1 and th (tyrosine hydroxylase, involved in dopamine biosynthesis) genes also show significant changes with transition from wnM to the vernal migratory LHS. Concurrent changes in the expression of genes associated with lipid metabolism and its transport also occur in the liver and flight muscles, respectively. Interestingly, there are clear differences in the behavioral and physiological phenotypes, and associated molecular changes, between the autumn and vernal migrations. In this review, we discuss seasonal changes in the behavior and physiology, and present molecular insights into the development of migratory phenotypes in latitudinal avian migrants, with special reference to Palearctic-Indian migratory buntings.

Impact of Light at Night Is Phase Dependent: A Study on Migratory Redheaded Bunting (Emberiza bruniceps)

Artificial light at night (LAN) alters the physiology and behavior of an organism; however, very little is known about phase-dependent effects of LAN, particularly, in night migratory songbirds. Therefore, in this study, we investigated whether the effects of LAN on daily activity and photoperiodic responses in the Palearctic Indian migratory songbird, redheaded buntings (Emberiza bruniceps), is dependent on the different phases of the night. Male buntings maintained under short photoperiod (8L:16D; L = 100 lux, D < 0.1 lux) in individual activity cages were exposed to LAN (2 lux) for 6 weeks either in 4 h bin given at the different phases of 16 h night (early, mid, or late at ZT 08–12, ZT 14–18, or ZT 20–24, respectively; n = 9 each group) or throughout 16 h night (all night light, n = 6, ZT 08–24, the time of lights ON was considered as Zeitgeber time 0, ZT 0). A group (n = 6) with no LAN served as control. The results showed that LAN at the different phases of night induced differential effects as shown by an intense activity during the night, altered melatonin and temperature rhythms, and showed an increase in body mass and body fattening, food intake, and gonadal size. Midnight light exposure has a greater impact on migration and reproduction linked phenotypes, which is similar to the ones that received light throughout the night. The highlights of this study are that (i) LAN impacts day-night activity behavior, (ii) its continuity with the day alters the perception of day length, (iii) birds showed differential sensitivity to LAN in a phase-dependent manner, (iv) the direction of placing LAN affects the daily responses, e.g., LAN in the early night was “accepted” as extended dusk but the late night was considered as early dawn, and (v) midnight LAN was most effective and induced similar responses as continuous LAN. Overall, LAN induces long day responses in short days and shows differential sensitivity of the different phases of the night toward the light. This information may be valuable in adopting a part-night lighting approach to help reduce the physiological burden, such as early migration and reproduction, of artificial lighting on the nocturnal migrants.

Daytime light spectrum affects photoperiodic induction of vernal response in obligate spring migrants

It is not well understood how the spectral composition (wavelength) of daylight that varies considerably during the day and seasons affects photoperiodic responses in a seasonal species. Here, we investigated the molecular underpinnings of wavelength-dependent photoperiodic induction in migratory redheaded buntings transferred to 13 h long days in neutral (white), 460 nm (blue), 500 nm (green) or 620 nm (red) wavelength that were compared with one another, and to short day controls for indices of the migratory (body fattening and weight gain, and Zugunruhe) and reproductive (testicular maturation) responses. Buntings showed wavelength-dependent photoperiodic response, with delayed Zugunruhe and slower testis maturation under 620 nm red light. Post-mortem comparison of gene expressions further revealed wavelength-dependence of the photoperiodic molecular response. Whereas there were higher retinal expressions of opn2 (rhodopsin) and opn5 (neuropsin) genes in red daylight, and of rhodopsin-like opsin (rh2) gene in green daylight, the hypothalamic opn2 mRNA levels were higher in blue daylight. Similarly, we found in birds under blue daylight an increased hypothalamic expression of genes involved in the photoperiodic induction (thyroid stimulating hormone subunit beta, tshb; eye absent 3, eya3; deiodinase type 2, dio2) and associated neural responses such as the calcium signaling (ATPase sarcoplasmic/endoplasmic reticulum Ca2+ transporting 2, atp2a2), dopamine biosynthesis (tyrosine hydroxylase, th) and neurogenesis (brain-derived neurotrophic factor, bdnf). These results demonstrate transcriptional changes in parallel to responses associated with migration and reproduction in buntings, and suggest a role of daylight spectrum in photoperiodic induction of the vernal response in obligate spring avian migrants.

Effect of T-photocycle and wavelength on photoperiodic induction in Indian weaver bird (Ploceus philippinus)

ABSTRACT Photoperiodic responses of a photosensitive species depend on the species-specific critical day length (CD), determined by the entrainment of the endogenous circadian rhythm of photoinducibility. We examined the entrainment pathway of the circadian photoperiodic system of photosensitive weaver birds. Birds were subjected to T = 22,24 and 26 h photocycles, with 11- and 12-h photophase to study the photoperiodic induction and synchronization of bird’s circadian system with reference of different light wavelengths. Two experiments were performed on adult male weaver birds. In experiment 1, birds were exposed to T = 22 and 26 h LD-cycles, having 11-h photophase with red and blue light wavelength, while in experiment 2, birds were exposed to T = 22, 24 and 26 h LD-cycles, having 12-h photophase with red and blue light wavelength. Observations on change in body mass and testis size were recorded at monthly intervals. There was a significant effect of red light wavelength on photoperiodic induction of testicular growth in the birds exposed to T = 22 h (11L:11D and 12L:10D) and T = 26 h (11L:15D and 12L:14D; both wavelength), but not when exposed to T = 24 h (12L:12D) photophase. Further, body mass regression was faster in birds of T = 26 h photophase (12L:14D) under blue wavelength. Thus, the above results are indicative of the influence of light wavelength on the endogenous programming that times seasonal responses in body fattening and testicular growth-regression of Indian weaver bird.

Illuminated night alters behaviour and negatively affects physiology and metabolism in diurnal zebra finches

Light at night (LAN) negatively impacts the behaviour and physiology; however, very little is known about molecular correlates of LAN-induced effects in diurnal animals. Here, we assessed LAN-induced effects on behaviour and physiology, and examined molecular changes in the liver of diurnal zebra finches (Taeniopygia guttata). Birds were exposed to dim LAN (dLAN: 12L = 150 lux: 12D = 5 lux), with controls on 12L (150 lux): 12D (0 lux). dLAN altered daily activity-rest and eating patterns, induced nocturnal eating and caused body fattening and weight gain, and reduced nocturnal melatonin levels. Concomitant increased nighttime glucose levels, decreased daytime thyroxine and triglycerides levels, and hepatic lipid accumulation suggested the impairment of metabolism under dLAN. Transcriptional assays evidenced dLAN-induced negative effects on metabolism in the liver, the site of metabolic homeostasis. Particularly, increased g6pc and foxo1 mRNA expressions suggested an enhanced gluconeogenesis, while increased egr1 and star expressions suggested enhanced cholesterol biosynthesis and lipid metabolism, respectively. Similarly, overexpressed sirt1 indicated protection from the metabolic damage due to elevated gluconeogenesis and cholesterol biosynthesis under dLAN. However, no effect on genes involved in lipogenesis (fasn) and insulin signalling pathway (socs3 and insig1) might indicate for the post transcriptional/post translational modification effects or the involvement of other genetic pathways in LAN-induced effects. We also found daily rhythm in the hepatic expression of selected clock and clock-controlled genes (per2, bmal1 and reverb-beta), with an elevated mesor and amplitude of per2 oscillation, suggesting a role of per2 in the liver metabolism. These results demonstrate dLAN-induced negative effects on the behaviour and physiology, and provide molecular insights into metabolic risks of the exposure to illuminated nights to diurnal animals including humans in an urban setting.

Transcriptional Signatures in Liver Reveal Metabolic Adaptations to Seasons in Migratory Blackheaded Buntings.

The molecular underpinnings of metabolic adaptation to seasons are poorly understood in long- distance migrants. We measured changes in physiology and performed de novo sequencing of RNA extracted from liver samples collected at 4-h intervals over a period of 24 h from a long-distance avian migrant, the blackheaded bunting (Emberiza melanocephala), during two states: photostimulated vernal migratory (M) state and photorefractory non-migratory (nM) state. The M state was differentiated from the nM state based on body fattening and weight gain, as well as on Zugunruhe, that is, nocturnal migratory restlessness in caged birds. We found that baseline blood glucose and triglyceride levels were significantly higher in the M state than the nM state; conversely, surface body temperature was higher in the nM state than the M state. In a total of 6 liver samples that were sequenced from each state, 11,246 genes were annotated, including 4448 genes that were cyclic over 24 h. We found 569 differentially expressed genes (DEGs) between the M and the nM state, and the M state showed 131 upregulated and 438 downregulated genes. These DEGs formed core gene hubs associated with specific biological processes in both the states. In addition, weighted gene coexpression network analysis revealed two discrete modules of coexpressed genes, with a significant difference in the expression pattern of metab olism-associated genes between M and nM states. These results demonstrate, for the first time, transcriptome-wide changes in the liver between two distinct physiological states and give molecular insights into seasonal metabolic adaptations in latitudinal migrants.

Annual life history-dependent gene expression in the hypothalamus and liver of a migratory songbird: insights into the molecular regulation of seasonal metabolism.

Birds seasonally switch from one life history state (LHS) to another to maximize their fitness. Accordingly, they exhibit distinct differences in their physiological and behavioral phenotypes between seasons. Possible molecular mechanisms underlying changes through the seasons have scarcely been examined in migratory birds. The present study measured key genes suggested to be involved in the metabolic regulation of 4 photoperiodically induced seasonal LHSs in a long-distance migratory songbird, the blackheaded bunting (Emberiza melanocephala). Buntings were held under short days (8 h light:16 h darkness, 8L:16D), during which they maintained the winter nonmigratory phenotype. Then they were exposed for several weeks to long days (13L:11D). Differences in the activity-rest pattern, body fattening and weight gain, testis size, organ (heart, intestine) weights, and blood glucose and triglyceride levels confirmed that buntings sequentially exhibited spring migration-linked premigratory, migratory, and postmigratory LHSs under long days. The mRNA levels of circadian genes involved in metabolism (Bmal1, Clock, Npas2, Rorα, and Rev-erbα) and of genes that encode for proteins/enzymes involved in the regulation of glucose (Sirt1, FoxO1, Glut1, and Pygl) and lipids (Hmg-CoA; Pparα, Pparγ; Fasn and Acaca) showed LHS-dependent changes in their light-dark expression patterns in the hypothalamus and liver. These initial results on genetic regulation of metabolism in a migratory species extend the idea that the transitions between LHSs in a seasonal species are accomplished by changes at multiple regulatory levels. Thus, these findings promise new insights into the mechanism(s) of adaptation to seasons in higher vertebrates.

Testes play a role in termination but not in initiation of the spring migration in the night-migratory blackheaded bunting

Day length regulates the development of seasonal phenotypes linked with both the spring migration (e.g., premigratory body fattening and intense nighttime restlessness, called Zugunruhe, in captive birds) and reproduction (e.g., gonadal growth and maturation). The apparent overlap in these photoinduced seasonal phenotypes could be taken to suggest that they are causally linked. The present study investigates this, using the night-migratory blackheaded bunting (Emberiza melanocephala). We continuously monitored activity of male buntings exposed for 19 weeks to short (8 h light : 16 h darkness; 8L : 16D) and long (16L : 8D) photoperiods. Long, but not short, days induced the spring migratory phenotype. Another experiment investigated the role of testes in spring migration by comparing the development of Zugunruhe between intact and castrated buntings held on natural day lengths, at Meerut (India, 29°01′N, 77°45′E) during the period from March to October, thus covering the times of both the spring and autumn migrations. Testes were not involved in induction of the migratory phenotype but probably influenced the end of the migratory season, since castrates delayed the termination of the Zugunruhe.

Temperature alters the photoperiodically controlled phenologies linked with migration and reproduction in a night-migratory songbird

We investigated the effects of temperature on photoperiodic induction of the phenologies linked with migration (body fattening and premigratory night-time restlessness, Zugunruhe) and reproduction (testicular maturation) in the migratory blackheaded bunting. Birds were exposed for four weeks to near-threshold photoperiods required to induce testicular growth (11.5 L:12.5 D and 12 L:12 D) or for 18 weeks to a long photoperiod (13 L:11 D) at 22°C or 27°C (low) and 35°C or 40°C (high) temperatures. A significant body fattening and half-maximal testicular growth occurred in birds under the 12 L, but not under the 11.5 L photoperiod. Further, one of six birds in both temperature groups on 11.5 L, and four and two of six birds, respectively, in low- and high-temperature groups on 12 L showed the Zugunruhe. Buntings on 13 L in both temperature groups showed complete growth-regression cycles in body fattening, Zugunruhe and testis maturation. In birds on 13 L, high temperature attenuated activity levels, delayed onset of Zugunruhe by about 12 days, reduced body fattening and slowed testicular maturation. The effect of temperature seems to be on the rate of photoperiodic induction rather than on the critical day length. It is suggested that a change in temperature could alter the timing of the development of phenologies linked with seasonal migration and reproduction in migratory songbirds.

Phase inversion of neural activity in the olfactory and visual systems of a night-migratory bird during migration

Olfactory and visual sensory mechanisms seem to play a critical role in migratory orientation and navigation. How these two mechanisms are functionally linked with other migratory processes is unknown. We investigated this, in relation to the profound behavioural shift that occurs during migration in the night-migratory blackheaded bunting (Emberiza melanocephala). Photosensitive unstimulated birds singly housed in activity cages were subjected to long days (LD 16/8). The activity of each bird was continuously monitored. Daily activity pattern defined the nonmigratory phase (no nocturnal activity) and migratory phase (intense nocturnal activity, Zugunruhe). Body mass and testis size were measured at the beginning and end of the experiment. Long days induced the migratory phenotype (body fattening and Zugunruhe) and testis maturation. The c-fos (Fos) immunoreactivity, as marker of the neural activity of the olfactory and visual subsystems, was measured at midday (8 h after lights-on) and midnight (4 h after lights-off) after the first seven long days (nonmigratory phase) and after seven nights of the Zugunruhe (migratory phase). In the nonmigratory phase, Fos-like immunoreactive (Fos-lir) cells in the olfactory and visual subsystems were high in the day and low at night. In the migratory phase, this was reversed; Fos-lir cells were high at night and low in the day. The phase inversion of neural activity in the olfactory and visual systems in parallel with the behavioral shift suggests a functional coupling between the systems governing migratory flight (expressed as Zugunruhe) and migratory orientation and navigation.

Short‐Days Induce Weight Loss in Siberian Hamsters Despite Overexpression of the Agouti‐Related Peptide Gene

Many vertebrates express profound annual cycles of body fattening, although it is not clear whether these represent differential activity of the central pathways known to mediate homeostatic control of food intake and energy expenditure, or whether the recent discovery of a major role for pars tuberalis-ependymal signalling points towards novel mechanisms. We examined this in the Siberian hamster (Phodopus sungorus) by using gene transfection to up-regulate a major orexigenic peptide, agouti-related peptide (AgRP), and then determined whether this increased anabolic drive could prevent the short-day induced winter catabolic state. Infusions of a recombinant adeno-associated virus encoding an AgRP construct into the hypothalamus of hamsters in the long-day obese phase of their seasonal cycle produced a 20% gain in body weight over 6 weeks compared to hamsters receiving a control reporter construct, reflecting a significant increase in food intake and a significant decrease in energy expenditure. However, all hamsters showed a significant, prolonged decrease in body weight when exposed to short photoperiods, despite the hamsters expressing the AgRP construct maintaining a higher food intake and lower energy expenditure relative to the control hamsters. Visualisation of the green fluorescent protein reporter and analysis of AgRP-immunoreactivity confirmed widespread expression of the construct in the hypothalamus, which was maintained for the 21-week duration of the study. In conclusion, the over-expression of AgRP in the hypothalamus produced a profoundly obese state but did not block the seasonal catabolic response, suggesting a separation of rheostatic mechanisms in seasonality from those maintaining homeostasis of energy metabolism.

Food deprivation during photosensitive and photorefractory life-history stages affects the reproductive cycle in the migratory Red-headed Bunting(Emberiza bruniceps)

In a seasonally breeding bird species, food deprivation affects reproduction. A key question is whether food shortage at any time in the year will affect reproduction even though the food supply subsequently becomes adequate. A prediction would be that a food supply that is of shorter duration than that optimally required during a life-history stage will have consequential effects on the succeeding life-history stages. Two experiments investigated this in gonadally regressed migratory Red-headed Bunting (Emberiza bruniceps) at two life-history stages: photosensitive and photorefractory. Experiment 1 employed two groups of photosensitive birds, and experiment 2 employed two groups of photorefractory birds. In both the experiments, birds were exposed for 8 weeks to a neutral day length at dim light intensity (12 h light: 12 h darkness, 12L:12D; L= approximately 5 lux, D=0 lux) with restricted feeding regimes (6 h food present: 18 h food absent, P:A 6:18 or P:A 12:12) and subsequently maintained for another 13 weeks at a highly stimulatory day length (16L:8D; L=400 lux; D=0 lux) with food ad libitum. We report that the pretreatment with restricted food cycles influenced the subsequent photoperiodic induction of reproductive (testis growth and molt) but not of metabolic (body fattening and mass gain) functions. The testicular response cycle under 16L:8D had a significantly lower amplitude in birds pretreated with P:A 6:18 than with P:A 12:12. Similarly, the recovery of photosensitivity was slower in photorefractory birds pretreated with P:A 6:18 than with P:A 12:12 food cycles. Overall, our findings show for the first time in a seasonally breeding vertebrate species that food deprivation during non-breeding periods of the annual cycle can affect reproductive functions later in the year.

Daily light regulates seasonal responses in the migratory male redheaded bunting (Emberiza bruniceps)

This study analyzed the role of day length in regulation of seasonal body fattening and testicular growth in a latitudinal Palaearctic-Indian migrant, the redheaded bunting (Emberiza bruniceps). When exposed to increasing photoperiods (hours of light: hours of darkness; 11.5L:12.5D, 12L:12D, 12.5L:11.5D, 13L:11D, 14L:10D, and 18L:6D) for 9-12 weeks, buntings responded in a photoperiod-dependent manner and underwent growth and regression cycle under photoperiods of > or =12 hr per day. Also, the response to a long photoperiod of birds that were held under natural photoperiods at 27 degrees N for 2 years was similar to those who arrived the same year from their breeding grounds ( approximately 40 degrees N), suggesting that the experience of higher amplitude day-night (light-dark, LD) cycles during migratory and breeding seasons were not critical for the subsequent response (initiation-termination-reinitiation) cycle. Another experiment examined entrainment of the circadian photoperiodic rhythm in buntings by subjecting them to T=24+/-2 hr LD-cycles with 8 hr photophase and to T=22 and 24 hr with 11 hr photophase. The results showed a reduction in critical day length under T=22 hr LD-cycle. In the last experiment, we constructed an action spectrum for photoperiodic induction by exposing birds for 4.5 weeks to 13L:11D of white (control), blue (450 nm), or red (640 nm) light at irradiances ranging from 0.028 to 1.4 W m(-2). The threshold light irradiance for photoinduction was about 10-fold higher for blue light, than for red and white lights. These results conclude that the daily light of the environment regulates the endogenous program that times seasonal responses in body fattening and testicular cycles of the redheaded bunting.

Wavelength Dependency of Light-Induced Effects on Photoperiodic Clock in the Migratory Blackheaded Bunting (Emberiza melanocephala)

The effects of light wavelength on photoperiodic clock were determined in the migratory male blackheaded bunting (Emberiza melanocephala). We constructed an action spectrum for photoperiodic induction (body fattening, gain in body mass, and gonadal recrudescence) by exposing birds for 4.5 weeks to 13 h light per day (L:D = 13:11 h) of white (control), blue (450 nm), or red (640 nm) color at irradiances ranging from 0.028 to 1.4 W m−2. The threshold light irradiance for photoinduction was about 10-fold higher for blue, compared to red and white light. Phase-dependent effects of light wavelength on the photoperiodic clock were further examined in the next two sets of skeleton photoperiods (SKPs). In the first set of SKPs, birds were exposed for four weeks to asymmetrical light periods (L:D:L:D = 6:6:1:11 h) at 0.25 ± 0.01 W m−2; two light periods applied were of the same (450 nm: blue:blue, B:B; 640 nm, red:red, R:R) or different (blue:red, B:R or red:blue, R:B) wavelengths, or of white:white (W:W, controls). Photoperiodic induction occurred under R:R and B:R, but not under B:B and R:B light conditions; the W:W condition induced an intermediate response. The second set of SKPs used symmetrical light periods (L:D:L:D = 1:11:1:11 h), and measured effects also on the activity rhythm. Birds were first exposed to one of the four SKPs (R:R, B:B, R:B, or B:R) for three weeks, subsequently were released into dim constant light (LLdim; ˜0.01 W m−2, the night light used in an L:D cycle) for two weeks, and then were returned to respective SKPs for another three weeks. Activity was greater in the R:R compared to B:B, and in B:R compared to R:B light condition. Zugunruhe (intense nighttime activity, indicating migratory restlessness in a caged situation) developed under the R:R and B:R, but not the B:B and R:B, light condition. Under LLdim, all birds free-ran with a period >24 h, the Zugunruhe had a circadian period longer than the daytime activity, and the re-entrainment to SKPs was influenced by the position of light periods relative to circadian phase of the activity rhythm. Photoperiodic induction at the end of 8 weeks was found in the R:R and B:R, but not in B:B, light conditions; in the R:B condition only one bird had initiated testes. Taken together, these results suggest that in the blackheaded bunting, the circadian photoperiodic clock is differentially responsive to light wavelengths; this responsiveness is phase-dependent, and the development of Zugunruhe reflects a true circadian function. Wavelength-dependent response of the photoperiodic clock could be part of an adaptive strategy in evolution of the seasonality in reproduction and migration among photoperiodic species under wild conditions.

Effects of Wavelength and Intensity of Light in Initiation of Body Fattening and Gonadal Growth in a Migratory Bunting Under Complete and Skeleton Photoperiods

Effects of Wavelength and Intensity of Light in Initiation of Body Fattening and Gonadal Growth in a Migratory Bunting Under Complete and Skeleton Photoperiods

Effects of wavelength and intensity of light in initiation of body fattening and gonadal growth in a migratory bunting under complete and skeleton photoperiods

A series of experiments investigated the effects of wavelength and intensity of light in initiation of body fattening (gain in body weight) and gonadal growth in migratory blackheaded bunting under complete and skeleton photoperiods. Using fluorescent light at an intensity ∼ 700 lx, the first experiment compared the inductiveness of a complete (13 h continuous light coupled with 11 h darkness; 13L:11D) and a skeleton (two light pulses of 6 and 1 h at 6 h apart; 6L:6D:1L:11D) photoperiod. Observations at the beginning and after 3, 8, 10, and 13 weeks of the treatment indicated that both photoperiods were fully inductive but that birds under 13L photoperiod fattened and lost body weight significantly earlier than birds under skeleton photoperiod. In the second experiment, bunting were subjected to 13L:11D ( L = 100 lx; D = 0 lx) of white, green (528 nm), and red (654 nm) light for a period of 5 weeks. Birds gained weight and testes grew in all groups except for an inconsistent fattening response in the white light group. The third experiment tested if the inductive effects of 1-h light pulse in a skeleton photoperiod were intensity dependent. Groups of bunting were exposed to 6L:6D:1L:11D (intensity of 1-h white light pulse = 2, 10, 50, or 100 lux) and examined at the beginning and after 3 and 8 weeks of the treatment. Photoinduction occurred at a slower rate and only at 50- and 100-lx intensities. The fourth experiment was similar to the third in design but it employed 1-h light pulse of two different wavelengths (green = 528 nm, and red = 654 nm) at 50- and 100-lx intensities. Birds fattened and testes grew only under red light. The last experiment varied the wavelength and intensity of the first (6 h) light pulse of the skeleton photoperiod (6L:6D:1L:11D): 6-h entraining light pulse of white, green (528 nm), or red (654 nm) colour at 10- or 50-lx intensity was used with 1-h inducing light pulse of white light at ∼ 700- lx intensity. Testes grew in all groups but significant fattening occurred only in birds entrained to 50-lx light intensity. These results indicate i) the dissociation of body weight and gonadal responses, in the sense that the timing of photostimulation and/or magnitude of photoperiod-induced body weight and testicular responses differed under various photoperiodic manipulations, and ii) the circadian processes involved in photoperiod-induced responses have differential spectral and intensity sensitivity.

Investigations of photoperiodically induced fattening in migratory blackheaded bunting (Emberiza melanocephala)(Ayes)

The body fattening and weight gain preceding vernal migration in birds is timed by a set of environmental factors of which daylength is predominant. However, the mechanism(s) by which these events is determined is poorly understood. Previous investigations on a photoperiodic migratory species, the blackheaded bunting (Emberiza melanocephala), indicate the involvement of a light‐sensitive circadian rhythm during initiation of fat deposition and body weight gain. This communication presents data from another set of experiments aimed to characterize further the mechanism(s) of fat deposition in the same species.Groups of photosensitive, unstimulated and stimulated birds were subjected to transfer and superimposition experiments for 30 days. While the former set included shifting of long‐day (LD) birds to DD, SD (short days), DD/LD and SD/LD, in the latter a 90‐minute bright light was superimposed at two different times of the day during the dim‐green lighted phase 15L:9D of varying intensity. Birds were weighed at the beginning and at the end of experiments. Those in transfer cycles were also weighed at 10‐day intervals. The results suggest that the premigratory body fattening and weight gain in blackheaded buntings is light dependent and timed by environmental daylength in accordance with the photosensitive endogenous circadian rhythm (ECR). They also show that the photoperiodic responses in birds in general are mediated by circadian rhythm(s).

Seasonal Variation of Reproduction and Fattening in Gulf Killifish from Brackish Mariculture Ponds

Seasonal changes in gonadal indices, plasma androgens in males, plasma estrogens and egg production in females, and body fat content in both sexes were monitored in gulf killifish, Fundulus grandis, reared in mariculture ponds. Changes in plasma androgens (males) and estrogens (females) were positively correlated with changes in gonadosomatic indices (GSI) of both sexes during most of the year. Seasonal variation in the GSIs of a nearby natural marsh population were similar, though not identical, to that of the pond-raised population. However several distinct differences were noted between these two populations: 1) spawning was prolonged in the mariculture population as compared with the wild population; 2) body fat content of the mariculture fish was significantly greater than wild fish during all seasons of the year; 3) a significant negative correlation between the seasonal cycles of body fattening and reproduction was observed in the mariculture fish but not in the wild fish. These data indicate that the mariculture environment and resultant high levels of body fattening supported prolonged spawning in this species.