Does overwinter temperature affect maternal body composition and egg traits in yellow perch Perca flavescens?

Female yellow perch Perca flavescens exposed to three overwinter temperature regimes (4, 8 and 13° C) for 150 days spawned in markedly different proportions upon spring warming (37% of females in 4° C v. 64 and 91% in 8 and 13° C treatments, respectively), but exhibited no differences in fecundity, egg size or egg lipid content. Females held at 4° C also exhibited less within‐clutch egg size variation than females held at 13° C. Moreover, eggs differed among temperature treatments in the overall proportions of 18 fatty acids, with the colder treatments resulting in potentially higher quality eggs containing more of the unsaturated fatty acids C16:1, C22:6‐n3 and C18:2 cis. Female somatic condition also varied with temperature. Maternal somatic growth and protein content increased while lipid content decreased in 13° C compared to the colder treatments. There were, however, no differences among treatments in the fatty acid composition of maternal muscle. These results suggest that the temperatures experienced during winter may be less influential to P. flavescens egg size or number, which may exhibit relatively little plasticity in this species, but can alter both the number of females that spawn and the overall composition of eggs and maternal somatic tissues, which may have implications for future reproductive success.     

Physiological flexibility: the key to success and survival for Antarctic fairy shrimps in highly fluctuating extreme environments

1. The anostracan fairy shrimp Branchinecta gaini inhabits one of the most hostile environments on earth, living in pools and lakes in Antarctica. Between January 2002 and January 2003 temperatures in two pools where B. gaini are extremely abundant on Adelaide Island ranged from ?18.6 to ?15.7 °C in winter, to 19.4 to 17.1 °C in summer, whilst air temperatures ranged from ?34 to 6.3 °C. 2. Branchinecta gaini survives winter as cysts, but endures large summer temperature fluctuations as adults. Cysts froze between ?24.4 and ?25.7 °C. In experiments adults survived 0–10 °C with no mortality for 1 week, 25 °C for nearly 48 h with 50% mortality, and at 32 °C complete mortality occurred in <1 h. 3. Oxygen consumption (M?O₂) in B. gaini approximately doubled for every 10 °C temperature rise (Q₁₀ = 2.04) up to 20 °C where it reached a peak. Females had, on average 19% higher M?O₂ than males. Females also had greater metabolic scopes, (maximum–minimum M?O₂ across temperatures was ×3.6 for females, ×3.1 for males). 4. Ventilation frequency increased linearly with temperature, and did not decline at 25 °C, indicating animals were ‘trying’ progressively harder to supply oxygen to tissues, and oxygen deficiency was the probable cause of death. Females had a higher ventilation frequency than males (8.6–17.1% higher) and they also exhibited greater scope to raise ventilation frequency (×2.4 for females versus ×1.5 for males). 5. Great metabolic flexibility allows B. gaini to exploit extreme, highly fluctuating environments, and larger ventilatory and respiratory scopes allow females to survive higher temperatures than males. Because of this flexibility their prospects for coping with physical environmental change are high.     

ECOTYPIC VARIATION IN PHYLLOPHORA PSEUDOCERANOIDES (RHODOPHYTA) ENSURES WINTER REPRODUCTION THROUGHOUT ITS GEOGRAPHIC RANGE1

Responses to temperature and daylength were determined in laboratory culture for isolates of the red alga Phyllophora pseudoceranoides (Gmelin) Newroth et A.R.A. Taylor from Nova Scotia, Iceland, Roscoff (France), and Helgoland (Germany). All isolates grew from 3° to 25° C and survived from -2° or 0° C to 27° C but not 30° C. Reproductive requirements differed between life history phases and isolates. Isolates from Helgoland and Roscoff formed sporangial sori at 3°-20° C, tetraspores at 3°-12° C, and procarps at 10°-20° C, irrespective of daylength. Spermatangia developed at 10°-23° C but only in long days. As the other European isolates, the isolate from Iceland formed tetrasporangia at 3°-12° C, but it had an additional requirement for short days. The Nova Scotian isolate formed sori at 10°-20° C and sporulated at 10°-18° C. When grown plants were transferred from noninductive to inductive conditions, sori were formed after 4 months and tetraspores developed and were shed (1-)3 months later. Procarps formed 1(-3) months after transfer. The phenology of P. pseudoceranoides was studied at Helgoland and Roscoff, where similar seasonal patterns were observed. Plants were perennial, forming new blades from October to June, which degenerated between August and February. In June, reproductive structures (sori, spermatangia, and procarps) started to appear on the new blades. From October to April, mature cystocarps were found. Mature tetrasporangia were observed only in February. The life history of P. pseudoceranoides is regulated by temperature and daylength. Differential effects on the different life history phases all serve to confine the production of spores (both carpospores and tetraspores) to the winter season. Differences in response between isolates from different geographic regions bring about the same effect: spores are shed only in winter. The nature of the geographic boundaries of P. pseudoceranoides is discussed.     

The effect of temperature on loose smut of wheat (Ustilago nuda)

In experiments made in controlled-temperature rooms on the wheat variety Kota, inoculated with two races of Ustilago nuda (Jens.) Rostr., maximum smut development occurred at 23 °C; at 20 ° and 15 °C the incidence and extent of smutting was reduced. An even greater reduction occurred when seedlings were grown at 6 °C for 2 months. No marked effect of temperature on mycelial development within the spike was observed until the ear emerged, when the mycelium in many of the ears of plants subjected to the lower temperatures failed to sporulate. The date of sowing in the field influenced disease development, presumably through the period of exposure to winter temperatures. Variety Little Club inoculated with race C₃ showed a similar but less marked response to cold-treatment.     

Effects of temperature and photoperiod on ovarian recrudescence in the cyprinid fish Mirogrex terrae-sanctae

Mirogrex terrae-sanctae is a commercial, sardine-like cyprinid fish in the Sea of Galilee. The fish spawns from December to the end of March. Females collected at the quiescent phase of the cycle (June) were exposed to four combinations of photoperiod and temperature for 3 months. The highest GSI (2.44 ± 0.22) was found in females exposed to simulated winter conditions (16°C, 9L) while the lowest GSI (0.65 ± 0.04) was found in fish exposed to simulated summer conditions (27°C, 14L). The GSI of fish maintained at the unnatural combination of 16°C, 14L (1.30 ± 0.09) did not differ from that of fish maintained at the other unnatural combination of 27°C, 9L (1.07 ± 0.11). However, histological examinations revealed that vitellogenic oocytes were present only in fish maintained at low temperature regimes, which implies that the growth of the ovarian mass in the 27°C, 9L group was due mainly to oogonial proliferation and that vitellogenesis was inhibited in Mirogrex at a high temperature. Both protein and calcium concentrations increased in the serum of fish following injections of oestradiol-17β. The increase of calcium was evident only in the non-filtrable (bound) fraction, suggesting that the total serum calcium above 13 mg% could be used as an indirect parameter for circulating vitelloprotein. Fish collected at the quiescent phase were injected with oestradiol (50 μg/fish c. 15 g body weight) and subsequently transferred to tanks maintained at either 12°, 20° or 27°C. The total serum calcium increased to 23.0 ± 1.75 mg% in the fish maintained for 8 days at 20°C. This was significantly higher than the concentration of calcium in the serum of fish maintained at either 12°C (7.8 ± 0.52) or 27°C(l1.8 ± 0.77). This implies that the temperature may affect ovarian recrudescence by modulating the response of the liver to oestrogen, allowing the oestrogen-induced vitellogenesis to be relatively high in the cool environment during winter.     

The influence of physiological age of Argas reflexus larvae (Acari: Argasidae) and of temperature and photoperiod on induction and duration of diapause

The occurrence of diapause and quiescence was investigated in Argas reflexus engorged larvae, nymphs I and nymphs II. For diapause experiments, larvae were maintained at five different locations: at constant 20°C long day (LD; 17 h light:7 h dark) or short day (SD; 10 h light:14 h dark), at two locations with natural photoperiod and temperature and at one location with natural photoperiod but constant 15°C. At 20°C, diapause incidence was low in physiologically young larvae, increased with larval age, and then decreased to zero in specimens of increased physiological age. This pattern, observed both at constant LD and SD, suggests that the propensity to diapause changes with the physiological age of the unfed larva. The duration of diapause decreased with increasing larval physiological age at all locations, resulting in a seasonally synchronized moulting pattern. The results suggest that A. reflexus larvae are photoperiodically sensitive both before and after feeding and that decreasing daylengths may be particularly strong inductive stimuli. The developmental zero and thermal constant of the larvae were determined as 13.24°C and 220 degree-days, respectively. Degree-day measurements revealed that larval A. reflexus may enter a diapause of different length when fed between August and December and kept at natural daylength. Development of engorged nymphs I and nymphs II, but not of larvae, was ultimatively restricted at a temperature of 37.5°C, but immediately resumed at 25°C, demonstrating the occurrence of quiescence at high temperatures. Similarly, at a low temperature of 15°C, many nymphs I and II did not develop within 58 months, but did so successfully after transfer to 25°C, without additional food intake. Received: 20 May 1997 / Accepted: 4 August 1997     

Biological and ecological studies on the sugar-beet fly,Pegomyia mixta Vill. (Diptera: Anthomyiidae) on sugar-beet in Egypt

The sugar-beet fly, Pegomyia mixta Vill., is the most serious insect pest affecting sugar-beet plantations in Egypt. This study wase carried out in field in the El-Nubaria region ofEl-Behare Governorate. Peak numbers of flies were taken in sweep nets in December. Development of the fly appeared to be restricted to the months between November and May. In the hot months, adults were most active early in the morning and late in the afternoon, but in the cold months the peak of activity occurred at about midday. The flies were generally found on the upper surfaces of the leaves at temperatures below 16 °C and on the lower surfaces at temperatures above 24 °C. Females were generally more numerous than males. The eggs were observed from the first week of November to the end of April; the population was three blotches of eggs and 17 larvae/20 plants. The highest infestation of this insect in El-Nubaria was recorded at the end of March in both seasons (10 blotches of eggs, 20 larvae, and five pupae, and 12 blotches of eggs, 22 larvae, 13 pupae, respectively). The eggs are deposited in groups (3–8 eggs). The larvae bore its blotch in the leaf, about 3–7 larvae may be found in one blotch.     

Maturity and growth of Paralonchurus brasiliensis females in southern Brazil (Teleostei, Perciformes, Sciaenidae)

Samples of Paralonchurus brasiliensis Steindachner, 1875 (627 females; no males) were taken monthly from June 1997 to May 1998 at the fishing pier of Cidreira – Rio Grande do Sul (30°08′S; 50°11′W) in southern Brazil. The fish were captured by sport fishermen using different sizes of hook‐and‐line. The reproductive period of P. brasiliensis extends from August to March, presenting greater activity from October to February. Females reach first maturation at 15.4 cm total length (TL). Of the 466 scales analyzed, one to six annuli were found. Growth annuli are formed between June and August, corresponding to the winter period (southern hemisphere), with the first visible annulus being formed when the fish are 1.5 years old. The length growth curve for P. brasiliensis females is described as: L_t = 20.0 (1 ? e^?0.564×t), TL in centimeters and age in years. The undersize asymptotic length may be a result of excessive fishing mortality on the studied population.     

Northern grass lizards (<Emphasis Type="Italic">Takydromus septentrionalis</Emphasis>) from different populations do not differ in thermal preference and thermal tolerance when acclimated under identical thermal conditions

We acclimated adults of Takydromus septentrionalis (northern grass lizard) from four localities (populations) under identical thermal conditions to examine whether local thermal conditions have a fixed influence on thermal preference and thermal tolerance in the species. Selected body temperature (Tsel), critical thermal minimum (CTMin), and critical thermal maximum (CTMax) did not differ between sexes and among localities in lizards kept under identical laboratory conditions for ∼5 months, and the interaction effects between sex and locality on these measures were not significant. Lizards acclimated to the three constant temperatures (20, 25, and 35°C) differed in Tsel, CTMin, and CTMax. Tsel, CTMin, and CTMax all shifted upward as acclimation temperature increased, with Tsel shifting from 32.0 to 34.1°C, CTMin from 4.9 to 8.0°C, and CTMax from 42.0 to 44.5°C at the change-over of acclimation temperature from 20 to 35°C. Lizards acclimated to the three constant temperatures also differed in the range of viable body temperatures; the range was widest in the 25°C treatment (38.1°C) and narrowest in the 35°C treatment (36.5°C), with the 20°C treatment in between (37.2°C). The results of this study show that local thermal conditions do not have a fixed influence on thermal preference and thermal tolerance in T. septentrionalis.     

The reproductive ecology of the invasive Asian paddle crab,Charybdis japonica (Brachyura: Portunidae), in northeastern New Zealand

The Asian paddle crab, Charybdis japonica, native to the northwest Pacific, is an invasive species that has established populations within northeastern New Zealand. Here, we provide a detailed examination of C. japonica's reproductive seasonality and gametogenesis, sizes at physiological maturity, and sex ratios outside its native range. Trapping in the Weiti River Estuary, Whangaparoa (36°38.4′S, 174°43.6′E) from February 2010 to May 2012 indicated a male‐biased population. Reproductive seasonality largely depended on female condition, since males contained ripe spermatozoa year‐round. Female gametogenesis began during late autumn, with oocyte development occurring throughout winter to early spring. Based on the presence of mature gametes during late winter and the appearance of gravid females in early November, mating began during early spring, with spawning in mid‐spring. Spawning coincided with a large drop in the gonad index, and occurred between sea surface temperatures of 17–22°C and 11.75–13.5 h day length. Females remained reproductive for up to 5 months of the year. Histological observation suggested that CW50 (the carapace width at which 50% of the population is mature) for females was 46.39 mm, while CW50 for males could not be determined due to a paucity of immature males. Histological analysis also suggested that females of C. japonica could produce store sperm and multiple broods annually.     

The effect of resistant cultivars of catch crops on the hatching of Heterodera schachtii

In laboratory tests with Heterodera schachtii, root leachates of nematode-resistant and susceptible cruciferous crops stimulated egg hatches of up to 77% and 95% respectively. In these tests very little hatch occurred at 5°C or 10°C, more at 15°C and most at 20°C. In field experiments in which these crops were sown in September 1982 and August 1983 into soil infested with H. schachtii, numbers of viable eggs had decreased only slightly under resistant crops 4–5 months later. This was possibly because the soil was too cold and dry to permit a large proportion to hatch.     

Thermal habitat of adult Atlantic salmon Salmo salar in a warming ocean

The year-round thermal habitat at sea for adult Atlantic salmon Salmo salar (n = 49) from northern Norway was investigated using archival tags over a 10 year study period. During their ocean feeding migration, the fish spent 90% of the time in waters with temperatures from 1.6–8.4°C. Daily mean temperatures ranged from −0.5 to 12.9°C, with daily temperature variation up to 9.6°C. Fish experienced the coldest water during winter (November–March) and the greatest thermal range during the first summer at sea (July–August). Trends in sea-surface temperatures influenced the thermal habitat of salmon during late summer and autumn (August–October), with fish experiencing warmer temperatures in warmer years. This pattern was absent during winter (November–March), when daily mean temperatures ranged from 3.4–5.0°C, in both colder and warmer years. The observations of a constant thermal habitat during winter in both warmer and colder years, may suggest that the ocean distribution of salmon is flexible and that individual migration routes could shift as a response to spatiotemporal alterations of favourable prey fields and ocean temperatures.     

Effects of temperature, light, storage conditions, sowing time, and burial depth on the seed germination of Cardiocrinum cordatum var. glehnii (Liliaceae)

In this study, we conducted experiments to accumulate practical information on the propagation and establishment of a population of Cardiocrinum cordatum var. glehnii by seed sowing. C. cordatum var. glehnii seeds require approximately 19 months from seed dispersal to cotyledon emergence in the field. However, the period from seed dispersal to radicle emergence was shortened to approximately 7–8 months by the temperature transition of 25/15°C (60 days) → 15/5°C (30 days) → 0°C (120 days) → 15/5°C (i.e., 15/5°C represents alternating temperature treatment wherein the seeds were placed at 15°C for 12 h during the day and then at 5°C for 12 h during the night). More than 90% of the seeds, which were stored dry at 5°C for 12 months and sown in pots in the field, showed cotyledon emergence, whereas in seeds stored dry at 25°C, dry at room temperature, and non-dry at room temperature, cotyledon emergence was decreased by less than 1%. More than 88% of the seeds that were stored dry at 5°C and sown in the field in October 2002 immediately after collecting, November, and from April to July 2003 showed cotyledon emergence in spring 2004. However, seeds sown in August, September, and October 2003 showed cotyledon emergences of 57.6%, 0%, and 0% in spring 2004, respectively. Seeds collected in October 2002 and sown until July 2003 in the field received adequate high temperature in summer, moderate temperature in autumn, and cold temperature in winter; therefore, the percentage of cotyledon emergence was high in spring 2004. On the other hand, seeds sown in August 2003 or later could not receive enough high temperature; thus, cotyledons emerged from only a few seeds.     

Geographic variations in life-history traits of the intertidal ocypodid crab Macrophthalmus banzai

Reproductive traits of the mud crab Macrophthalmus banzai were compared in populations from Okinawa (26°13′ N) and Amakusa (32°32′ N). Crabs matured earlier and at a smaller size in Okinawa than in Amakusa, but growth rate did not differ between the two localities. Population density was much higher in Okinawa. Breeding occurred in winter in Okinawa and in summer in Amakusa. In Okinawa, females produced one brood at over 1 year of age and then died. In Amakusa, females produced one brood at over 1 year of age, and one or two broods at around 2 years of age, and then died. Egg weight in the Okinawa population was almost twice that of the Amakusa population. Furthermore, brood weight was larger in Okinawa than in Amakusa. Longevity was 16–19 months in Okinawa and 28–30 months in Amakusa. At high latitudes, the winter low temperature blocks reproduction of M. banzai. At low temperature blocks reproduction of M. banzai. At low latitudes, their short lifespan prevents iterative breeding. Thus, the difference in breeding season greatly affects other life-history traits.     

Effects of temperature and food level on growth and development of a planktonic water mite

We analysed the relative effects of food availability and temperature on rates of growth and development of a predatory planktonic water mite, Piona exigua. Growth in length of mites fed Daphnia, Ceriodaphnia and Chydorus was analysed by Gompertz or von Bertalanffy curves; these curves were compared by parallel curve analysis. Growth rates of nymphs and adult female mites increased with temperature; the duration of the imagochrysalis stage decreased. Females grown at 10 °C were smaller at final size than females grown at 15 °C, 18 °C or 22 °C. Females reared at food levels of 15 or 30 prey l⁻¹ grew more slowly and were smaller than those provided with 60 or 120 prey l⁻¹. Nymphs grew more slowly when Daphnia were the only prey, than when smaller prey were available. Food level did not affect nymph growth at 10 °C or 15 °C, but growth at 18 °C or 22 °C may have been slowed at the lowest food levels. Synergistic effects of temperature and food level on nymph growth were apparent only from analysis of growth curves and not from stage duration data.     

Adaptation to habitat in Aquilegia species endemic to Sardinia (Italy): Seed dispersal,germination and persistence in the soil

Abstract

The autecology of the Sardinian endemics Aquilegia barbaricina Arrigoni et Nardi and A. nugorensis Arrigoni et Nardi were investigated. Peaks of anthesis and seed dispersal were recorded for five populations occurring in two distinct habitats, one riparian and one rupicolous. Germination tests were carried out on seed lots belonging to each population by sowing seeds at 10, 15, 20, 25 and 25/15°C. In addition, seeds were incubated for 2 months at either 25°C (summer), 5°C (winter) or 25°C for 2 months plus 2 months at 5°C (summer followed by winter–SW), and then moved to the germination temperatures. Embryo measurements were taken during pre-treatments and germination. Experimental seed burials were carried out for two populations of each species. Both species dispersed in summer. The population of A. nugorensis occurring on rocky outcrops differed in phenology from both the other A. nugorensis population from riparian vegetation and from A. barbaricina. Both species showed morphophysiological seed dormancy, with <50% germination under laboratory conditions. All riparian populations germinated only after the SW pre-treatment, while the rupicolous population germinated at 25°C, without any pre-treatment. Low germination percentages were observed in the experimental seed burials, suggesting the ability for both species to form a persistent soil seed bank.     

Comportamento Alla Germinazione Delle Cariossidi Delle Linee Primaverile e Invernale del Triticum Esaploide «Denti de Cani»

Abstract

The germination of spring and winter wheat lines of exaploid Triticum « Denti de Cani ». — The dormancy in the seeds of two lines of Triticum « Denti de Cani » (which is spontaneous in Sardinia), one with solid stem (CP line), a spring line, the other with hollow stem (CV line), an winter line, has been studied. Germination was carried out in the dark, in Petri dishes at the constant temperatures of 5°, 10°, 20°, 23°, 26°, 30° and 35°C, using full ripe seeds, and seeds in different stages of after-ripening up to one year of age. The increase in % germination, for increasing temperatures above 5°C, is clearly conditioned by the progress of after-ripening in the seeds. In fact it was seen that, in general for the two lines, percentages over 50% of seeds germinated at 3 days were reached: at 10° and 20° after 15 days from the full ripening; at 23°C after 30 days; at 26°C after 50 days; at 30°C after about 100 days and at 35°C only after about 4–5 months from the harvest. During the experiment at 5°C it was observed that, during the first year of life of seeds and especially in the CP line, this temperature produces a clear slowing down in germinations after first year from the ripening, only the CV seeds — not the CP which remain very much inhibited — reach germination values over 50% at 3 days. It has also been demonstrated that the CV are more sensitive than the CP, in the first initial period of after-ripening (15 and 30 days), to the non-inhibiting activity of low temperatures (5° and 10°C) and that, between these, the 10°C temperature promotes the germination more clearly than the 5°C temperature. The results obtained have shown that the dormancy wears off in the spring CP-line much more slowly than in the winter CV-line. The CP-seeds remain in a relative dormancy condition for a long time, which causes a significative delay in germination, up to 100 days from the full ripening stage.     

Reproductive parameters of clouded leopards (Neofelis nebulosa)

Reproductive data compiled from the International Clouded Leopard Studbook revealed that 75% of all litters were born to females between one and five years of age. Sixty-three percent of the males had sired litters by four years of age with reproduction declining after six years of age. Sixteen zoological institutions surveyed worldwide contributed estrous cycle data from 28 clouded leopards. Sexual maturity (age at first estrus) for these females ranged from 17 to 28 months of age with a mean estrous cycle length of 29.9 ± 13.8 days. The mean length of estrus was six days. Gestation length ranged from 85 to 121 days (X? = 93.4 ± 6.3). There was a significantly higher incidence of estrus in fall and winter compared with spring and summer over latitudes ranging from 36°–45° and 51°–55° (P < .005). Mating occurred in all months except June and October, and cubs were produced in all months except December. The highest frequency of mating occurred during the month of December corresponding with a birth peak in March.     

Isolation of yeasts from palm tissues damaged by the red palm weevil and their possible effect on the weevil overwintering

Rhynchophorus ferrugineus is a tropical pest of palms that has recently invaded Japan, where winter temperatures fall below 0°C. Because activities of the weevil at temperatures <13°C are extremely limited, it appears difficult for them to overwinter in Japan. However, the temperature of palm tissues damaged by this weevil has been observed to be higher than air temperature. Here, we looked for the cause of this temperature increase. First, we measured the temperature of damaged palm tissues and showed it to be between 30°C and 40°C, even in winter. Next, we isolated yeasts from the body of weevils and infested palm tissues and obtained 36 yeast strains, mostly Candida tropicalis and C. ethanolica. Then, we analyzed the soluble sugar composition in palm tissues and found that it included glucose, sucrose, and fructose. Because at least C. tropicalis can ferment some of these sugars, the temperature increase may be attributed to fermentation of microbes, including yeasts.     

SEASONAL CHANGE IN THE DISTRIBUTION OF CELL SIZE OF COCCONEIS SCUTELLUM VAR. ORNATA (BACILLARIOPHYCEAE) IN RELATION TO GROWTH AND SEXUAL REPRODUCTION1

Growth and sexual reproduction of the marine littoral diatom Cocconeis scutellum Ehrenb. var. ornata Grun. were investigated at 30 different combinations of temperature (5, 10, 14, 18, 22° C), irradiance (20, 60, 100 μE·m^?2·s^?1) and daylength (14:10 and 10:14 h LD cycle). Growth occurred at all combinations. The optimal growth was observed at 14–18° C, long daylength and highest-to-moderate irradiance, and at 18° C, short daylength and highest irradiance. Sexual reproduction on the other hand occurred between 5 and 18° C, and the optimal condition was 10–14° C and short daylength. Annual cyclic, and sesonal changes in the distribution of cell size (valve length) were observed in a field population. These changes were characterized by an annual minimum in mean cell size in autumn, an annual maximum in winter, a slight decrease from the mean in spring–middle summer, a rapid decrease from the mean in late summer–early autumn, and appearance of bimodal distribution of cell size in winter. These changes were caused by sexual reproduction in autumn, rapid growth in late summer–early autumn and slow growth in other seasons, and poor viability of small cells near the lower end of the size range.