大猿叶虫生活史的研究

大猿叶虫Colaphellus bowringi是江西山区十字花科蔬菜上的重要食叶害虫,以成虫在土中越夏和越冬。由于成虫滞育期的差异,该虫的化性显示了明显的种内变异。有些个体隔年繁殖;有些个体是一化性的,仅在春季或秋季繁殖1代;有些个体是二化性的,春季和秋季各繁殖1代;有些个体是多化性的,春季1代,秋季2～3代。因此,大猿叶虫在田间一年可发生4代。在春季,滞育成虫于2月底至4月初陆续出土繁殖;在秋季,滞育成虫于8月中旬至10月初陆续出土繁殖。春季羽化的成虫于4月底至6月上旬陆续入土越夏,秋季羽化的成虫于9月中旬至12月底陆续入土越冬。大猿叶虫一生能交配多次,大多数雌虫的产卵期超过1个月,最长达67天。平均每雌产卵量：春季世代为644粒,秋季世代为963粒,最高达1.950粒。各虫态的发育历期：在15～30℃间,卵为13.78～3.14天,幼虫为22.83～6.95天,蛹为12.10～3.18天。发育阈值温度：卵为10.7℃,幼虫为8.8℃,蛹为9.6℃。非滞育成虫的寿命约为1～2个月,滞育成虫的寿命为5～38个月。滞育成虫均入土蛰伏,在土中的蛰伏深度为9～31 cm 。     

Ecophysiological attributes of adult overwintering in insects: insights from a field study of the nut weevil,Curculio nucum

Abstract Diapausing insect species have evolved a great diversity of life cycles, although overwintering occurs at a single development stage within most species. Understanding why diapause has evolved towards a given life stage requires investigation of both the ecological and physiological attributes. Notably, it is suggested that adult overwintering is more energy‐demanding than larval overwintering but it brings fitness gains by allowing adults to be synchronized with their seasonal requisites through an early spring emergence. This hypothesis is tested in field conditions in the nut weevil Curculio nucum, whose life cycle comprises an obligate 2‐year, nonfeeding underground phase, including a larval, followed by an adult, overwintering. In this species, adult wintering leads to an early spring emergence; at first glance, however, this does not enhance synchronization between weevils and their host because adults emerge more than 1 month before starting to breed. It is suggested that adult overwintering ultimately evolved in response to the phenology of the host, by allowing females to oviposit in nuts before their full sclerotinization. Adult overwintering appears to be costly because adults postpone reproduction for 1 year, incur a significant weight loss and require feeding before egg laying. Surprisingly, lipids are unaffected during diapause, lipogenesis even being likely in the summer metamorphosis. These results suggest that the lipids involved in egg production may entirely come from the larval stages, whereas the other nutrients are acquired through adult feeding but this remains to be tested.     

Reproductive behavior in captive California condors (Gymnogyps californianus)

From the fall of 1987 to the spring of 1994, data on reproductive behavior were collected on seven different pairs of California condors (Gymnogyps californianus) housed at the San Diego Wild Animal Park. Behavioral data were collected from videotaped activities in their flight pens. Behaviors for this report include male and female displays and incomplete and complete copulations. Data on the time it took for females to recycle and lay another egg after egg removal were acquired from the California Condor Studbook. Male display rates were significantly higher than female rates throughout the breeding season. Males began displaying 52–132 days before the start of copulations. Copulations slowly increased from Day -51 and peaked ∼ Day -8, prior to egg laying. Female display rates were significantly more frequent at the end of the season after the last egg was laid. An average of 26.8 (±3.1 SEM) copulations occurred prior to egg laying. Copulations occurred throughout the day, with the higher mean percentages occurring between 0700 and 0900 hours. After egg removal, the interval between laying of the first and second eggs and between the second and third eggs had a mean of 34.1 (±10.1 SD) days. © 1996 Wiley-Liss, Inc.     

Interaction between flight,reproductive development and oviposition in the pine weevil Hylobius abietis

• 1 The development of reproductive and flight capacity of pine weevils Hylobius abietis during the spring and their dispersal to, and subsequent development at, new clearfell oviposition sites comprise key phases in their life cycle in managed forests. At an old clearfell site where autumn‐emerging weevils had overwintered, weevils were trapped as they re‐emerged in the spring and tested for their ability to fly and then dissected to determine the degree of wing muscle and egg development.
• 2 Re‐emerging weevils were most abundant in pine growing at the edge of the clearfell and, over most of the trapping period (April to June), their capacity for flight (proportion flying and wing muscle width) was more advanced than in weevils from the clearfell itself, with a similar trend in the degree of reproductive development (proportion with mature eggs and egg volume).
• 3 In weevils from the clearfell, flight capacity and reproductive development increased concurrently to a peak around mid‐May. In weevils from pine, wing muscles were already well developed at the start of trapping, although few of them flew. Their more advanced development was attributed to the increased opportunities for maturation feeding after emergence in the previous autumn.
• 4 In the spring, weevils reached the canopy of trees for maturation feeding by walking and, to a lesser extent, by flight. Weevils dispersed by flight to oviposition sites in mid‐May when most of them were reproductively mature. After arrival, flight ability and wing muscle size declined rapidly but egg production was maintained until most weevils had stopped flying. When wing muscles reached their minimum size, there was a marked decline in egg size, suggesting that wing muscle breakdown is important in maintaining egg production at oviposition sites. Prospects for further wing muscle and reproductive development are discussed.

     

Life cycle of three stonefly species (Plecoptera) from an Apenninic stream (Italy) with the description of the nymph of Nemoura hesperiae

The life cycles of Isoperla grammatica, Amphinemura sulcicollis and Nemoura hesperiae are studied in a Northern Italy stream, sited in the Apennines. The three species show a univoltine life cycle and their development coincides approximately in the same period of the year, although the one of I. grammatica is longer. Possible egg dormancy could exist in the A. sulcicollis and N. hesperiae life cycles, but this needs to be proved. The three taxa show a spring flight period in the study area. Growth is almost constant through the life cycle, except in A. sulcicollis in which an increase is observed at the end of the development. N. hesperiae exhibits a faster growth than the other species. In addition, in this paper, the nymph of N. hesperiae is described and designed for the first time.     

Timing matters: species‐specific interactions between spawning time,substrate quality,and recruitment success in three salmonid species

Substratum quality and oxygen supply to the interstitial zone are crucial for the reproductive success of salmonid fishes. At present, degradation of spawning grounds due to fine sediment deposition and colmation are recognized as main factors for reproductive failure. In addition, changes in water temperatures due to climate change, damming, and cooling water inlets are predicted to reduce hatching success. We tested the hypothesis that the biological effects of habitat degradation depend strongly on the species‐specific spawning seasons and life‐history strategies (e.g., fall‐ vs. spring‐spawners, migratory vs. resident species) and assessed temperature as an important species‐specific factor for hatching success within river substratum. We studied the species‐specific differences in their responses to such disturbances using egg‐to‐fry survival of Danube Salmon (Hucho hucho), resident brown trout (Salmo trutta fario), and migratory brown trout (Salmo trutta lacustris) as biological endpoint. The egg incubation and hatching success of the salmonids and their dependence on temperature and stream substratum quality were compared. Hatching rates of Danube salmon were lower than of brown trout, probably due to higher oxygen demands and increased interstitial respiration in spring. Increases in maximum water temperature reduced hatching rates of resident and migratory brown trout (both fall‐spawners) but were positively correlated with hatching rates of Danube salmon (a spring‐spawner). Significantly longer incubation periods of resident and migratory brown trout coincided with relatively low stream substratum quality at the end of the egg incubation. Danube salmon seem to avoid low oxygen concentrations in the hyporheic zone by faster egg development favored by higher water temperatures. Consequently, the prediction of effects of temperature changes and altered stream substratum properties on gravel‐spawning fishes and biological communities should consider the observed species‐specific variances in life‐history strategies to increase conservation success.     

Mating and reproduction of predaceous diving beetles,Dytiscus sharpi,observed under artificial breeding conditions

Mating season and embryonic development of the predaceous diving beetles, Dytiscus sharpi, (Coleoptera; Dytiscidae) were observed under artificial breeding conditions. Female and male adult insects started mating from November to March and gave first instar larvae mainly in April. When the mating was artificially delayed until February, first instar larvae appeared from the end of March to the middle of May. I also investigated the effects of temperature on larval development. Apparent hatchability of eggs was not affected by high temperature, however, their normal development after hatching was significantly interfered. Most of the first instar larvae kept at 20-25 degrees C from before hatching died within one day after hatching. By contrast, juveniles kept outdoors (7.0-20.9 degrees C) could develop at least until second instar larvae. Temperature >23 degrees C after hatching had no effects on larval development. From these observations, it was concluded that the reproduction strategy of Dytiscus sharpi, i.e. mating in late autumn and hatching in early spring would be the reasonable results of adaptation to the warm habitats where they are collected.     

THE DEVELOPMENT OF BEHAVIOUR IN ELEUTHERODACTYLUS MARTINICENSIS (AMPHIBIA, ANURA)

In the genus Eleutherodactylus , development is wholly embryonic. Among other features of ontogeny which are thereby modified is the development of behaviour. Within the egg envelope, the embryo is largely still, but wriggles from time to time. These movements represent all that remains of the axial swimming behaviour of larval Amphibia. The embryo develops normally when removed from the envelopes, and its reactions to light touch can then be studied from day to day. In E. martinicensis , which develops from first cleavage to hatching in 13-14 days, the first movement of the legs is seen during trunk wriggles at 7 1/2 days, but a day later they can move independently. The first reaction of the leg is a slow withdrawal; at 9 1/2 days flexion is followed by a vigorous extension. By then, reflexes affecting all four limbs can be elicited.
Towards the end of embryonic life, limb movements can be maintained in continuous cycles, and the animal can then swim. When it is allowed normally to emerge from the envolopes, the juvenile toad, smaller than a house fly, can jump a distance of over a foot. These observations are discussed in relation to the ontogeny of behaviour in other vertebrates, particularly in regard to the views of G. E. Coghill.     

Oögenesis defects in the ecd-1 mutant of Drosophila melanogaster, deficient in ecdysteroid at high temperature

Oögenesis defects observed in ecd-1 females depend on the stage at which the shift from the permissive temperature (20°C) to the non-permissive temperature (28.5°C) is performed. Pupae shifting earlier than 48 hours after puparium formation have underdeveloped ovaries and die at emergence. In ecd-1 females shifted 2 days or more after puparium formation, egg chambers develop normally up to the early vitellogenic stage, and then degenerate—no mature oöcytes are formed. When the shift is performed after maturation of the females at 20°C, their egg-laying rate decreases slowly after the shift; the eggs display abnormal chorions and fail to develop into larvae. By shifting the ecd-1 females back to 20°C, a complete reversal of abnormalities of eggshell formation and egg development may be observed. The temperature-sensitive periods related to these different defects were determined. For egg development, the period extends from 2 days to 1 day before laying. For eggshell formation, the beginning and end of the temperature-sensitive period were not separated—both were located around 24 hours before laying. For the block of vitellogenesis, we concluded that this is probably due to an inhibition of yolk uptake. This inhibition would be effective only in follicles younger than the previtellogenic stage 7 when submitted to the non-permissive temperature.     

Notes on the life history of the clam shrimp,Eulimnadia texana

Several life history measures (growth rate, egg production, molt frequency, age at maturity and lifespan) were measured on several clam shrimp hermaphrodites (Eulimnadia texana Packard) grown in a laboratory setting under optimal growth conditions. Growth rates were high early in life, and then dropped dramatically when egg production began (day 5–6). Early egg production was low, and increased until approximately day 7, after which production leveled off for several days. Reproductive senescence was noted after day 17, with clutch sizes continuously dropping until death. Average molts per day was approximately 1.1, and molting seemed to be more closely associated with egg production than with growth. Growth and egg production were negatively correlated, indicating a possible trade-off between these two traits. No other trade-offs were detected. These shrimp show typical early-colonist life history traits, displaying high initial growth, early reproduction at a high rate, and then early senescence and death. This revised version was published online in August 2006 with corrections to the Cover Date.     

Early life history strategies of notothenioids at South Georgia

Antarctic notothenioid early life history strategies are examined in general and then for common species at South Georgia. Channichthyids, bathydraconids, artedidraconids and some nototheniids have large eggs 3·0–4·9 mm whereas other nototheniids and arpagiferids have smaller eggs 1·6–2·7 mm. At South Georgia the larvae of species with large eggs hatched between August (late winter) and late November (late spring) at 11–16 mm standard length ( L _s). Larvae of species with small eggs hatched mainly during October and December at 4.5–9 mm L _s. Most of the larvae of all species attain urostyle flexion between October and January, and develop to the end of the larval stage between November and May. The duration of the larval stage varies from 2 months in species with smaller larvae to 6 months for some of the species with larger larvae. Two nototheniid species develop to the early juvenile stage before a channichthyid and a bathydraconid that hatch around 2 months earlier. During their first winter, the early-juveniles of most species with large eggs are pelagic, whereas those of species with small eggs may be pelagic or demersal. Four groups of strategies are proposed based on egg size and the winter ecotype of the early-juvenile stage.     

Testis development and testosterone secretion in captive European ground squirrels before, during, and after hibernation

In some hibernating species, an extended euthermic period before spring emergence has been reported during which testicular maturation occurred. In this study, we investigated whether male European ground squirrels Spermophilus citellus (Linnaeus, 1766) develop scrotal testes before or after the termination of hibernation. The course of testis development and testosterone concentrations were examined in young-of-year male ground squirrels (n = 4) before, during, and after their first hibernation. In the environmental chamber, all animals showed regular hibernation patterns with decreasing torpor bout lengths prior to the end of hibernation. Four weeks before hibernation ended, testosterone levels were elevated during spontaneous arousals. Testosterone concentrations peaked during the first 2 months post heterothermy and decreased thereafter. In 2 experimental males, testes descended from the inguinal region into the scrotum shortly before the end of heterothermy. The state of testis development and testosterone secretion corresponded to that found in semi-free-living males at spring emergence. Testis size increased in active animals to maximum widths during weeks 3 and 4 post heterothermy. Scrotal skin pigmentation was absent until heterothermy ended and developed during the first month thereafter. The data suggest that male S. citellus do not need a pre-emergence euthermic period for reproductive development.     

Daily egg production ofTemora longicornis (Copepoda,Calanoida) during winter and early spring in the Marsdiep (Southern North Sea)

Mature females of the copepodTemora longicornis from the Marsdiep tidal inlet were incubated at ambient water temperature during the winter of 1989/1990 to estimate their egg production during 24 hours. Throughout winter this production remained at a level of 0 to 10 eggs per female. In March a steep rise to 60 to 80 eggs coincided with an equally steep rise in chlorophyll-a from 4 to 20 g.l^–1. Excess food given during incubation tended to increase the observed production rate only in October at relatively high temperature. Additive effects of rising food level and temperature on daily egg production were found in spring. In March and April 1988 and 1990 the observed maximum daily egg production per female increased from about 20 at 5°C and 5 g.l^–1 chlorophyll to 70 at almost 10°C and 20 g.l^–1. The egg production in winter gives a prebloom potential to maintain a stock of premature stages ready to utilize the algal spring bloom. Early spawning before April can also contribute to the development of the abundance peak of adults in May–June. Due to growth rates increasing with temperature in the months March and April, most of the adults at their peak density may have hatched from eggs before the period of maximum egg production rate at the end of April. The exceptionally high production rate observed at the end of March may be related to a developingPhaeocystis bloom.     

Laboratory studies on egg and faecal pellet production of Centropages typicus: effect of age, effect of temperature, individual variability

The rates of egg production of individually reared females ofCentropager typicus have been observed from the moulting C5-adultto death under constant conditions of food (10 000 cells ml^–1of Hymenomonas elongara) and temperature (15 and 20°C).Preliminaiy experiments showed that virgin females could produceunfecunded eggs, but that the distinctive spines of the eggsof C.typicus were only obtained after fecundation occurred.High and continuous egg production needs several matings, andthe following experiments were run with a couple. In all experiments,both males and females fed and produced faecal pellets. Thefemales show a high variability in spawning life periods, inspawning rhythms, in daily rates and, consequently, in cumulatednumbers of eggs spawned during their life. The highest dailyrates of egg production were obtained in the first half of thefemale's life, and were as high at 20°C as at 15°C.Nevertheless, the females survived longer at 15°C than at20°C, and finally produced 47.6% more eggs during theirwhole life at 15°C than at 20°C, which can be partiallydue to the difference in size (almost 10% bigger at 15°Cthan at 20°C). The production of faecal pellets occurredduring the whole of life and was more regular than egg production,with a maximum daily rate in the first part of life, and showeda decrease until death. The decline in daily egg productionwith age was parallel to the decrease in daily faecal pelletproduction. The daily production of faecal pellets was higherat 15°C than at 20°C. At 20°C, a good correlationwas observed between the total egg production per female andthe total faecal pellet production per female. The individual-basedegg production rates of our experiments have been compared withrates obtained in different conditions by several authors, andwere lower than in other published studies, probably due tothe food offered. We emphasize the difficulties of comparingresults which have been obtained for the entire life of thefemales with those based on daily egg production. The totalegg production of a female is an important parameter in regardto the population d because it fixes the maximal rate of losses(mortality plus dispersion) for which a population can continueto develop.     

INTERACTIVE EFFECTS OF OFFSPRING SIZE AND TIMING OF REPRODUCTION ON OFFSPRING REPRODUCTION: EXPERIMENTAL,MATERNAL, AND QUANTITATIVE GENETIC ASPECTS

We demonstrate that egg size in side-blotched lizards is heritable (parent-offspring regressions) and thus will respond to natural selection. Because our estimate of heritability is derived from free-ranging lizards, it is useful for predicting evolutionary response to selection in wild populations. Moreover, our estimate for the heritability of egg size is not likely to be confounded by nongenetic maternal effects that might arise from egg size per se because we estimate a significant parent-offspring correlation for egg size in the face of dramatic experimental manipulation of yolk volume of the egg. Furthermore, we also demonstrate a significant correlation between egg size of the female parent and clutch size of her offspring. Because this correlation is not related to experimentally induced maternal effects, we suggest that it is indicative of a genetic correlation between egg size and clutch size. We synthesize our results from genetic analyses of the trade-off between egg size and clutch size with previously published experiments that document the mechanistic basis of this trade-off. Experimental manipulation of yolk volume has no effect on offspring reproductive traits such as egg size, clutch size, size at maturity, or oviposition date. However, egg size was related to offspring survival during adult phases of the life history. We partitioned survival of offspring during the adult phase of the life history into (1) survival of offspring from winter emergence to the production of the first clutch (i.e., the vitellogenic phase of the first clutch), and (2) survival of the offspring from the production of the first clutch to the end of the reproductive season. Offspring from the first clutch of the reproductive season in the previous year had higher survival during vitellogenesis of their first clutch if these offspring came from small eggs. We did not observe selection during these prelaying phases of adulthood for offspring from later clutches. However, we did find that later clutch offspring from large eggs had the highest survival over the first season of reproduction. The differences in selection on adult survival arising from maternal effects would reinforce previously documented selection that favors the production of small offspring early in the season and large offspring later in the season—a seasonal shift in maternal provisioning. We also report on a significant parent-offspring correlation in lay date and thus significant heritable variation in lay date. We can rule out the possibility of yolk volume as a confounding maternal effect—experimental manipulation of yolk volume has no effect on lay date of offspring. However, we cannot distinguish between genetic effects (i.e., heritable) and nongenetic maternal effects acting on lay date that arise from the maternal trait lay date per se (or other unidentified maternal traits). Nevertheless, we demonstrate how the timing of female reproduction (e.g., date of oviposition and date of hatching) affect reproductive attributes of offspring. Notably, we find that date of hatching has effects on body size at maturity and fecundity of offspring from later clutches. We did not detect comparable effects of lay date on offspring from the first clutch.     

Age, growth and reproduction of the cyprinid Rutilus lemmingii (Steindachner, 1866) in the River Huebra, Central Spain

Age, growth and reproduction of the R. lemmingii population of the River Huebra, Duero basin are analysed. Females dominated older age classes and lived up to 6 years (5+) while males only reached 5 years (4+). The growing season extended from April to September; growth rates were similar for both sexes. 0+ fish condition increased during their first summer and spring. In older fish, condition cycle was related to gonad development and showed some differences between sexes. The number of females reaching maturity at age 1 + almost doubled that of males; females also matured at smaller size. Both fecundity and egg size increased with female length: mean egg counts varied between 974 for 1 + individuals and an estimated 10491 for 5+ fish. Eggs were produced as a single batch, but were released fractionally during April and May.     

The effects of constant and changing tempertures on the development of eggs of the freshwater snail Lymnaea auricularia (L.)

The rate of development of Lymnaea auricularia eggs was studied at various constant temperatures between 10° and 36°C. Development was accelerated as the temperature increased and at 36°C the eggs failed to develop. Spring eggs showed differences in their rate of development when compared with summer eggs when measured at similar tempertures.

Both spring and summer eggs were more than 90% fertile. Hatching success was high at temperatures between 10° and 30° (100%–82/9%); while at 34°C it was reduced to 60.6% for spring eggs. It was above 87% at temperatures between 10° and 34°C but it dropped to 62.3% at 36°C for summer eggs.

In one regularly changing temperature experiment a significant acceleration (P < .05) was found. In two others there was no significant difference beween predicted and observed egg durations. In one suddenly changing temperature regime (1 day at 20°, 1 day at 30° and so on) a huge retardation of development was found. In the other suddenly changing experiment (1 day at 15°, 1 day at 25°) no significant difference was found.

The exposure of eggs to extreme temperature (4°C, freezing and 4°C caused a retardation in the race of subsequent development of eggs at 25°C.     

Population dynamics, growth and production of Sigara selecta (Fieber, 1848) (Hemiptera, Corixidae) in a Mediterranean hypersaline stream

1. This is the first study on the life cycle, growth and production of Sigara selecta, a Palearctic corixid species typical of brackish and saline waters, at the warmest limit of its European distributional range. The study combines field and laboratory approaches. 2. The S. selecta population studied was multivoltine, producing four asynchronous cohorts from early spring to December and overwintering in the adult state. Development time from egg to first adult ranged from 2 to 3 months. A minimum temperature threshold of 10 °C and diel amplitude of ≥10 °C were observed for reproduction and oviposition. 3. Maximum density and biomass were reached in mid spring and early autumn. The sex ratio was unbalanced, females dominating during most of the year, except in spring, when the sex ratio was balanced or dominated by males during the first adult emergence. 4. Laboratory rearing experiments at constant temperatures (18, 22 and 26 °C) pointed to a significant effect of temperature on egg development and nymphal growth. In the range of temperatures tested, both egg and nymphal instar duration decreased with increasing temperature. Mean nymphal development time varied from 43 days at 26 °C to 71 days at 18 °C, with a mean of 57 days. Survivorship was independent of temperature. 5. A reduction in nymphal and adult length was observed with increasing temperature. 6. Growth rates decreased with increasing body mass and increased as temperature increased. The first nymphal instar had the highest length increments and growth rates in all temperature treatments. 7. Satisfactory agreement was found between the field and laboratory degree‐days required for complete development from egg to first adult. At constant and variable thermal regimes, degree‐days decreased with increasing temperature. 8. Rate of growth in the field could be predicted with reasonable accuracy from a simple model obtained as a function of body mass. The model explained 67% of the variability in growth rates. 9. Annual production and production/biomass ratio (P/B) of S. selecta estimated by the Instantaneous Growth method were 1.28 g m^?2 year^?1 and 13.71, respectively. Spring and autumn cohorts contributed 32% and 54%, respectively, of total annual production. Maximum production corresponded to intermediate temperature periods, although summer production may have been underestimated because of the longer sampling interval relative to cohort interval production. The Size Frequency method underestimated production by at least 18% with respect to the Instantaneous Growth method.     

The effect of environmental temperature on the production of ultrasounds by isolated non-handled albino mouse pups

Albino mouse pups develop homoiothermy during the first 19 days of their life and show three distinct stages during this period. From day 1 to day 6 they are almost completely poikilothermic at low ambient temperatures; from day 7 to day 14 or 15 is a transitional period during which they develop homoiothermy at medium temperatures (e.g. 22°C); from day 15 or 16 to day 19 or 20 is the final stage when homoiothermy at very low ambient temperatures is developed. These changes are very closely correlated with changes in the ultrasounds produced by isolated pups within the 19 day period. Thus their acoustic response to cold is weak from day 1 to day 5 or 6 and very strong from day 6 or 7 to day 12 or 13, on exposure even to medium temperatures. From day 14 onwards the pups progressively stop producing ultrasounds depending on the ambient temperatures, and there is complete cessation of calling after day 19 or 20 even on exposure to temperatures as low as 2°C. In general, decreasing the ambient temperatures increases the rate and intensity of ultrasounds produced but very low ambient temperatures have an inhibiting effect and at 2–3°C breathing and calling cease completely after a time which varies with age of the pups.     

Endocrine correlates of parental care in an Antarctic winter breeding seabird, the emperor penguin, Aptenodytes forsteri

Plasma levels of luteinizing hormone (LH) and prolactin associated with parental behavior were measured in the Antarctic winter breeding emperor penguin, Aptenodytes forsteri. Males exclusively incubate the egg while females exclusively brood the nonhomeothermic young; both sexes alternate in rearing the homeothermic young. Birds were sampled on arrival from the sea through egg laying, incubation, and brooding. All parent birds lost their chicks at the end of the brooding period due to harsh weather but sampling continued. In females, LH titers dropped after egg laying but levels were restored when the birds returned from the sea to brood the chicks and were not depressed by high prolactin levels. Plasma prolactin remained low in males captured on arrival and kept until the free-living males finished incubation. In breeders, prolactin secretion increased during the prelaying period when day length decreased. Prolactin levels stayed elevated in males during incubation and in brooding females returning after a 2-month absence at sea. Prolactin values were higher in brooding females than in males ending incubation or returning in late brooding. These levels did not drop after chick loss, and the sexual difference in prolactin values was maintained after breeding failure. In emperor penguins, increased prolactin secretion appears to be triggered around the time of egg laying and continues, driven by an endogenous mechanism, through incubation and brooding until rearing is completed. Prolactin secretion independent of external stimuli may have evolved in pelagic seabirds to maintain parental care despite long absences at sea from the breeding colony.