Cell-specific and temporal aspects of gene expression in the chicken oviduct at different stages of the laying cycle

Egg formation and embryonic development occur as the yolk passes through the magnum, isthmus, and shell gland of the oviduct before oviposition in hens. The present study identified candidate genes associated with secretory function of the chicken oviduct after ovulation and contributing to egg formation and oviposition. Hens (n = 5 per time point) were euthanized to recover the reproductive tract when the egg was in the magnum (3 h after ovulation) and the shell gland (20 h after ovulation). Total RNA was extracted from each segment of the oviducts and subjected to Affymetrix chicken GeneChip analysis. Quantitative PCR and in situ hybridization analyses of selected genes confirmed the validity of the gene expression patterns detected using microarray analysis. In particular, ACP1, CALB1, CYP26A1, PENK, RCAN1 and SPP1 expression increased significantly in the shell gland between 3 h and 20 h postovulation, whereas only RCNA1 expression increased significantly in the magnum between 3 h and 20 h postovulation. Results of the high-throughput analysis revealed cell-specific and temporal changes in gene expression in the oviduct at 3 h and 20 h postovulation in laying hens provide novel insight into changes at the molecular and cellular levels of candidate genes related to formation of the egg and oviposition.     

Testosterone immunization blocks the ovulatory process in laying hens without affecting ovarian follicular development

The role of testosterone in the ovulatory process in hens has been largely neglected. The aim of the present study was to evaluate if testosterone plays an important role on the ovulatory process in laying hens. The effect of active and passive immunization against testosterone on ovarian follicular development and oviposition was studied. Egg laying percentage was evaluated in hens actively immunized against testosterone-BSA (T-AI; n = 6) or BSA (BSA-AI; n = 6). Oviposition was reduced as antibody titer increased in T-AI hens (r = -0.67; P < 0.01). Ovarian structures were assessed in three animals from each group. Follicles reached preovulatory size in both groups, nonetheless, in T-AI hens follicles at different stages of regression indicated that ovulation was blocked by treatment. In the remaining animals, preovulatory concentrations of progesterone and testosterone were determined. A preovulatory surge release of progesterone, preceded by a testosterone peak, was observed in the BSA-AI group (P < 0.05). In contrast, progesterone in T-AI animals remained at basal concentrations. Whereas, testosterone concentrations were significantly greater in T-AI as compared with BSA-AI animals (P < 0.05). Finally, to study the effect of passive immunization on oviposition, hens were passively immunized (PI) on four occasions, on alternate days with anti-T serum (T-PI; n = 10) or anti-BSA serum (BSA-PI; n = 8). During the 13-day period that preceded treatment, oviposition averaged 94.1%. Forty-eight hours after the first immunization, no egg was laid by 8 out of the 10 T-PI hens. During the 10 days following the first passive immunization, there was a reduction in the laying percentage that was significantly greater in T-PI hens (reduction of 52% in T-PI versus 29% in P-BSA, P < 0.01). In summary, these studies show that testosterone immunization hampers egg-laying without affecting ovarian follicular development, suggesting that testosterone has an important role in the ovulatory process in laying hens.     

Developmental arrest during embryonic development of the common chameleon (Chamaeleo chamaeleon) in Spain

Embryonic development of the common chameleon, Chamaeleo chamaeleon, was monitored from oviposition to hatching at a field site in southwestern Spain and in the laboratory under five experimental temperature regimes. Embryos were diapausing gastrulae at the time of oviposition; developmental arrest in the field continued as cold torpor during winter. Postarrest development in the field commenced in April, and hatching occurred in August, for a total incubation period of 10.5 mo. In the laboratory, one group of eggs was incubated at a constant warm (26 degrees C) temperature. The remaining treatments simulated field conditions and consisted of initial periods of warm temperature of 0, 27, 46, and 71 d, a subsequent 4-mo period of cold winter (16 degrees C) temperature, and a final period of warm (26 degrees C) temperature. Embryos in the constant warm temperature treatment were in diapause an average of 3 mo, with clutch means ranging from 2 to 4 mo. Hatching among clutches occurred over 2 mo. In contrast, for field and experimental eggs that experienced cold winter conditions, hatching within treatments occurred over 2-14 d; "winter" conditions synchronized development. The length of time between the end of cold conditions and hatching did not differ among treatments; development thus resumed as soon as temperature was suitable regardless of the initial period of warm temperature. Diapause in nature thus insures that embryos remain gastrulae after oviposition despite nest temperatures that may be warm enough to support development.     

Prostaglandin production by the largest preovulatory follicles in the domestic hen (Gallus domesticus)

An injection of 5 micrograms of gonadotropin-releasing hormone (GnRH) into hens 8 h prior to oviposition advanced the expected time of oviposition by approximately 1 h. The plasma concentration of progesterone increased approximately 1 h earlier in GnRH-injected hens in comparison to saline-injected hens. The plasma concentration of 13,14-dihydro-15-keto-prostaglandin F2 alpha (PGFM) increased significantly (p less than 0.05) at the time of oviposition in both the GnRH- and saline-injected hens. Significantly (p less than 0.05) greater concentrations of prostaglandin F2 alpha (PGF2 alpha) were assayed in media containing the largest preovulatory follicles collected at oviposition than in media containing the second and fifth largest preovulatory follicles collected at the same time. No prostaglandin was detected in media containing small, nonhierarchial follicles. The concentration of PGF2 alpha in media containing granulosa cells from the largest preovulatory follicle was significantly greater (p less than 0.05) than in media containing 4 times as many theca cells. Ovine luteinizing hormone (oLH) alone or in combination with arachidonic acid had no effect on PGF2 alpha output from granulosa cells collected 6 h before oviposition, whereas A23187 caused a small stimulation of PGF2 alpha output. However, treating cells first with oLH and then with A23187 stimulated a 15- to 20-fold increase in PGF2 alpha. None of these stimuli enhanced the already high output of PGF2 alpha when added to incubations of granulosa cells collected within 5 min after oviposition. These data suggest that the granulosa cells of the largest preovulatory follicle are the major intraovarian source of prostaglandin and that production of PGF2 alpha is associated with the preovulatory surges of gonadotropins and steroid hormones preceding oviposition.     

Thermal effects on reptile reproduction: adaptation and phenotypic plasticity in a montane lizard

Interspecific comparisons suggest a strong association between cool climates and viviparity in reptiles. However, intraspecific comparisons, which provide an opportunity to identify causal pathways and to distinguish facultative (phenotypically plastic) effects from canalized (genetically fixed) responses, are lacking. We documented the reproductive traits in an alpine oviparous lizard, and manipulated thermal regimes of gravid females and their eggs to identify proximate causes of life‐history variation. Embryonic development at oviposition was more advanced in eggs laid by females from high‐elevation populations than in eggs produced by females from lower elevations. In the laboratory, experimentally imposed low maternal body temperatures delayed oviposition and resulted in more advanced embryonic development at oviposition. Warm conditions both in utero and in the nest increased hatching success and offspring body size. Our intraspecific comparisons support the hypothesis that viviparity has evolved in cold‐climate squamates because of the direct fitness advantages that warm temperatures provide developing offspring. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 642–655.     

Effects of early prenatal exposure to 2,3,7,8-tetrachlorodibenzo-p-dioxin (TCDD) on postnatal reproduction in the laying hen (Gallus gallus)

This study demonstrates the long-term effects of very early embryonic exposure to a single dose of 2,3,7,8-tetrachlorodibenzo-p-dioxin (TCDD) (0, 10 and 20 ng/egg), administered before the beginning of embryonic development, on growth and reproductive performance in laying hens. Hatchability and body weight gain from 11 weeks onwards were significantly depressed in 20 ng treated hens. All hens started laying egg at around the same age and the laying performance of TCDD-treated hens was normal. No disturbances in the age-related pattern and concentrations of oestradiol, LH or FSH in plasma could be found but mean progesterone concentrations were significantly lower in 20 ng treated hens. Moreover, follicular distribution was changed with less small white follicles and smaller yellow follicles, which probably resulted in the lower egg weight of the 20 ng treated hens. At 43 weeks of age, hens treated in ovo with TCDD showed a retained right oviduct, mostly filled with clear fluid. From these results, it seems that in ovo exposure to TCDD interferes in the right oviduct regression during embryonic development and induces some changes in follicular distribution but without impairment of reproductive performance in the adult laying hen.     

Egg-laying behaviour and nest-site selection of domestic hens kept in small floor-pens

The development of egg-laying behaviour and nest-site choice is described in seven flocks of White Leghorn hens housed in small floor-pens. Special interest was directed towards social dominance and the effect of the presence of a cock on the hen's nest-site choice. In the weeks before the hens came into lay, they frequently examined the nests. Nest-entries only occurred some days before, or in the hours preceding, the first oviposition. Some hens were very conservative in their nest-site choice within clutches as well as between clutches. A number of hens only persevered in their nest-site choice within clutches, while a third category did not show any consistency in their choices. Individual hens were consistent in using either the floor or a nest-box as a nesting site.No relationship was found between social status and nest-site choice, nor between social status and floor-laying. Some evidence was provided that the presence of a cock could reduce the number of floor-eggs in a flock provided with roll-away boxes.It is postulated that the nest-examinations in the weeks before laying may serve for the selection of a suitable nest-site.     

Evidence for a physiological role of prostaglandins in oviposition by the hen

A single oral dose of indomethacin (25 mg) given 2--4 h before expected oviposition delayed oviposition by about 12 h and suppressed plasma PGE levels by about 90%. Intrauterine injection of PGE-1 (0.2--0.4 microgram/hen) to 12 hens pretreated with 25 mg indomethacin induced oviposition in all birds in about 7 1/2 min. Passive immunization with goat antiserum to PGE-1 delayed oviposition (69%) in 4 hens. It is concluded that prostaglandins play a functional role in oviposition.     

Embryonic growth and mobilization of energy and material during incubation in the checkered keelback snake, Xenochrophis piscator

We collected 20 checkered keelback snakes (Xenochrophis piscator) to study embryonic growth and mobilization of energy and material during incubation. Females laid eggs between late May and late June. The eggs were incubated at 27 degrees C (+/-0.3). One egg from each clutch was dissected at five-day intervals starting at oviposition. The mean incubation length at 27 degrees C was 48.9 days. We identified three phases of embryonic growth or yolk depletion in X. piscator. Phase 1, between oviposition and Day 20, was one of minimal transfer of energy and material from yolk to embryo. Phase 2, between Day 20 and Day 39-40, was characterized by increasingly rapid embryonic growth or yolk depletion. Phase 3, between Day 39-40 and hatching, was characterized by reduced embryonic growth or yolk depletion. Approximately 71% of dry mass, 53% of non-polar lipids and 66% of energy were transferred from the egg contents to the hatchling during incubation. Our data confirm that oviposition is not timed to coincide with the onset of rapid embryonic growth in oviparous squamate reptiles. The greater conversion efficiencies of energy and material from egg to hatchling in snakes can be attributed to their lower energetic costs of embryonic development and greater residual yolk sizes.     

Mortality of Apatania fimbriata (Insecta: Trichoptera) during embryonic, larval and adult life stages

1. Egg masses, oviposition site preferences, and abiotic and biotic factors causing mortality during embryonic, larval and adult life stages of Apatania fimbriata were studied. Laboratory investigations provided information on the temperature dependence of embryonic development, measured as an increase in egg volume.
2. A. fimbriata laid hemispherical egg masses, consisting of a transparent matrix containing a mean of 208 eggs. Egg masses were laid on stones situated just above the water surface in dark cavities in the stream bank.
3. Two hundred egg masses were mapped and individually monitored during embryonic development. There was no significant correlation between mortality during embryonic development and any of the abiotic parameters measured. First-instar larvae of Osmylus fulvicephalus consumed developing eggs, and chironomids preyed on newly hatched larvae.
4. A mean of seventy-two females emerged per metre of stream. Mortality during the 1993/94 life cycle was measured as a percentage of the potential number of eggs laid. Female mortality between emergence and oviposition was ≈ 80%. Eight per cent of individuals were lost during embryonic development. Larval mortality to emergence in 1994 was 11.3%. This indicates that the terrestrial life stage is probably decisive in the regulation of A. fimbriata populations.
5. Duration of embryogenesis at constant temperatures (4–20 °C) in the laboratory was described best by a negative exponential function. This species is cold stenothermal and there was no hatching success at 20 °C.
6. Egg volumes during embryonic development increased sigmoidally over time.     

Altitudinal variation in life cycle syndromes of California populations of the grasshopper,Melanoplus sanguinipes (F.)

Summary Life cycles of California populations of the grasshopper, Melanoplus sanguinipes, varied along an altitudinal gradient. Temperature records indicate a longer season at low altitude on the coast, based on computation of degree days available for development, even though summer air temperatures are cooler than at high altitude; this is a result of warm soil temperatures. At high and low altitudes there was a high proportion of diapause eggs oviposited, while intermediate proportions of diapause eggs occurred at mid altitudes. The low altitude, and especially sea level, populations diapaused at all stages of embryonic development, while at high altitudes most diapause occurred in the late stages just before hatch. Diapause was more intense at high altitudes. One result of diapause differences was delayed hatching in the sea level population. Nymphal development and development of adults to age at first reproduction were both accelerated at high altitude relative to sea level. At lower temperatures (27° C) there was a tendency for short days to accelerate development of sea level nymphs, but not high altitude nymphs. In both sea level and high altitude grasshoppers, short days accelerated maturation of adults to onset of oviposition at warm temperature (33° C); there was little reproduction at 27° C. Population differences for all traits studied appear to be largely genetic with some maternal effects possible. We interpret diapause variation at low and mid altitudes to be responses to environmental uncertainty and variations in development rates to be adaptations to prevailing season lengths.     

Mortality of Apatania fimbriata (Insecta: Trichoptera) during embryonic, larval and adult life stages

1. Egg masses, oviposition site preferences, and abiotic and biotic factors causing mortality during embryonic, larval and adult life stages of Apatania fimbriata were studied. Laboratory investigations provided information on the temperature dependence of embryonic development, measured as an increase in egg volume.
2. A. fimbriata laid hemispherical egg masses, consisting of a transparent matrix containing a mean of 208 eggs. Egg masses were laid on stones situated just above the water surface in dark cavities in the stream bank.
3. Two hundred egg masses were mapped and individually monitored during embryonic development. There was no significant correlation between mortality during embryonic development and any of the abiotic parameters measured. First-instar larvae of Osmylus fulvicephalus consumed developing eggs, and chironomids preyed on newly hatched larvae.
4. A mean of seventy-two females emerged per metre of stream. Mortality during the 1993/94 life cycle was measured as a percentage of the potential number of eggs laid. Female mortality between emergence and oviposition was ≈ 80%. Eight per cent of individuals were lost during embryonic development. Larval mortality to emergence in 1994 was 11.3%. This indicates that the terrestrial life stage is probably decisive in the regulation of A. fimbriata populations.
5. Duration of embryogenesis at constant temperatures (4–20 °C) in the laboratory was described best by a negative exponential function. This species is cold stenothermal and there was no hatching success at 20 °C.
6. Egg volumes during embryonic development increased sigmoidally over time.     

Progesterone injection and egg production in turkey hens

An arrest in laying associated with either a polyovarian follicle (POF) or a polycystic ovarian follicle (PCOF) syndrome has been reported in turkey hens photostimulated at an early age with a constant-light photoperiod. Hens expressing the POF or PCOF syndrome had stopped laying for several weeks, but the ovary contained an increased number of mature-size and larger follicles (POF hens), which were cystic (PCOF) in some of the hens. Hens with the POF or PCOF syndrome had plasma progesterone (P(4)) concentrations that were relatively high and without surges. We hypothesized that high plasma P(4) concentrations may block ovulatory surges of LH but not the growth or maintenance of hierarchical follicles leading to development of the POF or PCOF syndrome in turkey hens. In the first six studies, hens were photostimulated with either a 14L:10D or a 24L:0D photoperiod and, after laying for 1-38 wk, were then injected daily for up to 14 days with P(4) (up to 1.50 mg kg(-1) day(-1)) and necropsied. At all ages, the oviposition rate was reduced at a P(4) dosage of 0.17 mg kg(-1) day(-1). With dosages of 0.33 mg kg(-1) day(-1) or greater, however, ovipositions stopped in most hens within approximately 2 days. For hens laying for less than 15 wk, oviductal weight and number of hierarchical follicles of P(4)-injected hens were not different from control vehicle-injected hens, but the numbers of mature, cystic, and atretic follicles were increased. For hens laying for 38 wk, when treated with P(4), oviductal weight and number of hierarchical follicles decreased, but number of atretic follicles increased. No effect of photoperiod was found on egg production, oviductal weight, or follicle number, and none of the hens developed POF or PCOF syndrome in these experiments. Two additional experiments were conducted with hens early in the reproductive period that had been photostimulated with 14L:10D or 24L:0D and injected with P(4) (0.33 mg kg(-1) day(-1)) for 10 or 12 days but not necropsied until 3 wk after the last injection. Most of the hens photostimulated with the 24L:0D photoperiod and injected with P(4), and a few of the hens photostimulated with the 14L:10D photoperiod and injected with P(4), had developed the PCOF syndrome when necropsied. The hens with the PCOF syndrome had high levels of P(4) when necropsied. From these studies, we concluded that the PCOF syndrome can be induced early in the reproduction period by photostimulating turkey hens with a 24L: 0D photoperiod, injecting them for 10 to 12 days with P(4) at a dosage of 0.33 mg kg(-1) day(-1), and then waiting 3 wk for the PCOF syndrome to develop.     

The isoenzyme composition of liver pyruvate kinase from hens in ontogeny]

Only one isozyme M2 of pyruvate kinase was found in the liver of hens at all stages of embryonic and postembryonic development. No analogue to isozyme L from the liver of mammals was found. During embryogenesis and postnatal life, isozyme M2 is presented by two forms which differ in pI values. Throughout embryonic and postembryonic development, pyruvate kinase is presented by two forms which differ in their substrate affinity.     

Evolution of viviparity: variation between two sceloporine lizards in the ability to extend egg retention

The evolutionary transition between oviparity and viviparity in squamate reptiles presumably occurs via a gradual increase in the duration of egg retention, the production of thinner eggshells, and increases in the vascularity of maternal and embryonic tissues. The 'ease' of this transition may differ among taxa. For example, in the genus Sceloporus , the scalaris species group contains both oviparous and viviparous species, and female Sceloporus scalaris can extend egg retention facultatively in response to the absence of a suitable site for oviposition without impairing embryonic development. In contrast, the undulatus species group contains only oviparous species, and, while female Sceloporus virgatus can extend egg retention, doing so retards embryonic development. I tested several hypotheses that would explain the greater ability of 5. scalaris than S. virgatus to extend egg retention. In this study, female S. scalaris retained eggs for 19 d without affecting the mortality of embryos, total developmental time, or dry mass of hatchlings. In contrast, when female S. virgatus retained eggs for 18 d, embryos had very high mortality and eggs took significantly longer to hatch than control (non-retained) eggs, although the dry mass of hatchlings was not affected. The ability of S. scalaris females to retain eggs with little negative effect on embryonic development was associated with relatively large chorioallantois, relatively thin eggshells, and relatively small clutch masses. These observations suggest that phylogenetic differences in the ability to extend egg retention may facilitate or constrain the evolution of viviparity in some lineages.     

Effect of the ovulatory cycle on oviductal sperm storage in the domestic fowl

Anovulatory domestic hens (pregnant mare serum-treated) and normally cyclic domestic hens were artificially inseminated with 0.034 ml of pooled semen. A subsequent microscopic assessment of the uterovaginal sperm storage glands on days 1, 3, 6, 9 and 12 post-insemination indicated that the sperm glands emptied over time in the normally cyclic hens, but not in the anovulatory hens. The data suggest that events associated with ovulation and/or oviposition are important to the sperm gland emptying process.     

多胚发育寄生蜂腰带长体茧蜂在寄主亚洲玉米螟幼虫体内的发育

多胚发育的幼虫内寄生蜂腰带长体茧蜂Macrocentrus cingulum的卵、胚胎和幼虫在寄主亚洲玉米螟Ostrinia furnacalis幼虫血腔内发育,通常1枚卵可以分裂增殖为数百只胚胎。本文通过定时解剖寄生的寄主幼虫,初步了解了腰带长体茧蜂多胚的形成过程及其在寄主体内的发育情况。结果表明：以4龄末期亚洲玉米螟幼虫为寄主时,寄生蜂卵产入寄主体内10 min左右开始卵裂,1天左右,初级胚胎从卵壳中被释放出来。之后胚胎在胚外膜内持续分裂产生大量二级胚胎形成桑葚胚。寄生后3天左右,二级胚胎从胚外膜中被释放出来,进入胚胎发育阶段。寄生后6天左右,胚胎进入胚带形成阶段。寄生后8天左右,胚带伸长,头尾形成。寄生后9天左右,身体分节完成,部分幼虫孵化,蜕去胚外膜。寄生后13天左右,蜂幼虫从寄主体内啮出。胚胎在发育初期体积变化不大,但从胚带形成开始,体积则迅速增大。腰带长体茧蜂与另一多胚发育寄生蜂佛州点缘跳小蜂Copidosoma floridanum在胚胎发育进程上明显不同,体现了它们对各自寄主的适应。     

Embryonic growth and mobilization of energy and material in oviposited eggs of the red-necked keelback snake, Rhabdophis tigrinus lateralis

We used the red-necked keelback (Rhabdophis tigrinus lateralis) as a model animal to study embryonic growth and mobilization of energy and material in oviposited snake eggs. Females (N=12) laid eggs between late May and early June. Eggs were incubated at 30 (+/-0.3) degrees C. One egg from each clutch was dissected at five-day intervals starting at oviposition. Incubation length averaged 27.9 days. Three phases of embryonic growth or yolk depletion could be detected in this study. The first phase, between oviposition and Day 10, was one of minimal transfer of energy and material from yolk to embryo. The second phase, between Day 10 and Day 22-23, was characterized by increasingly rapid embryonic growth and yolk depletion. The third phase, between Day 22-23 and hatching, was characterized by a gradual reduction in embryonic growth and yolk depletion. Approximately 73.6% of dry mass, 50.0% of non-polar lipids and 57.8% of energy were transferred from egg to embryo during incubation. Embryos withdrew mineral from the eggshell mainly during the last quarter of incubation. Our data show that oviposition does not coincide with the onset of rapid embryonic growth in oviparous species of squamate reptiles that are positioned midway within the oviparity-viviparity continuum, and that the greater conversion efficiencies of energy and material from egg to hatchling in snakes can be mainly attributed to their lower energetic costs of embryonic development and greater residual yolk sizes.     

The effect of indomethacin on ovarian prostaglandin release in hens

Indomethacin, an inhibitor of prostaglandin (PG) synthetase, will block uterine muscle electromyographic activity (EMG activity) and oviposition at a midsequence oviposition and ovulation in domestic hens, but does not block the increase in EMG activity associated with the first ovulation of a sequence. To assess the potential relationship between prostaglandin release from the ovarian follicles and EMG activity in egg-laying hens, we determined the concentrations of PGF2 alpha, 13,14-dihydro-15-keto-PGF2 alpha (PGFM), and PGE2 in brachial, ovarian follicular and uterine venous plasma and tissues in relation to uterine muscle EMG activity at the first ovulation and at a midsequence oviposition. The concentrations were measured after an i.m. injection (25 mg/hen) of indomethacin. In control hens sampled hourly, beginning 4 h before the peak of EMG activity at the first ovulation of a sequence, there was a sharp increase (p less than 0.05) in concentrations of PGF2 alpha and PGFM in brachial vein plasma coincident with the increase (p less than 0.05) in uterine EMG activity. Hens pretreated with indomethacin also had increased plasma PGF2 alpha and PGFM levels (p less than 0.05) in brachial vein plasma and increased uterine EMG activity (p less than 0.05) at this time. Indomethacin treatment lowered but did not eliminate mean levels of PGF2 alpha in the venous effluent from the largest preovulatory follicle at the first ovulation (36.0 +/- 9.9 ng/ml vs. 14.4 +/- 1.8 ng/ml).(ABSTRACT TRUNCATED AT 250 WORDS)     

Influence of oviposition date and temperature upon embryonic development in Somatochlora alpestris and S. arctica (Odonata: Corduliidae)

In two bog-dwelling dragonflies, Somatochlora alpestris and S. arctica , the influence of oviposition date and temperature upon duration of embryonic development was studied. Egg diapause was facultative. With advancing season, the proportion of diapause eggs increases from 0 to 37% in S. alpestris and from 0 to 18% in S. arctica. Eggs needed at least 17 to 38 days for development. Hatching curves were temperature-independent in nearly all experiments but developmental rate increased at higher ambient temperature. In S. arctica , responses of developmental rate to temperature differed in eggs laid on different dates. In S. alpestris , duration of egg development decreased as season progressed. The duration of egg development of non-diapause eggs and proportion of diapause eggs in S. alpestris and possibly in S. arctica may also both be a function of female age at the time of oviposition. The ecological significance of the different development patterns is discussed.