Endogenous regulation of environmentally induced sexuality in a rotifer: a multigenerational parental effect induced by fertilisation

SUMMARY 1. Sexual reproduction in the heterogonic life cycle of many rotifers occurs when amictic females, which produce diploid eggs developing parthenogenetically into females, are environmentally induced to produce mictic females. Mictic females produce haploid eggs which develop parthenogenetically into males or, if fertilised, into resting eggs – encysted embryos which develop into amictic females after an obligatory diapause. 2. A Florida strain of Brachionus calyciflorus was used to test the prediction that amictic females hatching from resting eggs (Generation 1), and those from the next few parthenogenetic generations, have a lower propensity to produce mictic daughters in response to crowding than those from later parthenogenetic generations. In 10 replicate clones, populations initiated by amictic females from generations 1, 5, 8, 12 and 18 were exposed to a standardised crowding stimulus, and the proportion of mictic females in the populations was determined. These proportions varied significantly across generations and clones. They were very low in the early generations and gradually increased to a mean of about 0.5 at Generation 12. 3. The mechanism for the transgenerational plasticity in response to crowding is not known. One possibility is that resting eggs contain an agent from their fertilised mictic mother's yolk gland that prevents development into mictic females and is transmitted in increasingly low concentrations through successive parthenogenetic generations of amictic females. 4. This parental effect may contribute to clonal fitness by ensuring that a clone developing from a resting egg will attain a higher population size through female parthenogenesis before maximising its commitment to sexual reproduction, even in the presence of a crowding stimulus from a high population density of other clones. Therefore, the number of resting eggs to which a clone contributes its genes should be maximised. 5. The clonal variation in propensity to produce mictic females in this strain indicates genetic variation in the trade‐off between maximising population growth via female parthenogenesis and increasing the probability of producing at least some resting eggs before local extinction from the plankton.     

Induction of mictic females in the rotifer Brachionus: oocytes of amictic females respond individually to population-density signal only during oogenesis shortly before oviposition

1. One at a time during the reproductive period of amictic females, oocytes fill with yolk and undergo a mitotic maturation division (oogenesis), are oviposited as single cells, and then develop parthenogenetically into females. Sexual reproduction in Brachionus and several other genera is initiated when amictic females are crowded and oviposit some eggs induced to differentiate into mictic females. Mictic females produce haploid eggs that can develop parthenogentically into males or be fertilised and develop into diapausing embryos called resting eggs. 2. This study examines the time when oocytes in amictic females respond to maternal population density. Is the fate of all oocytes in the germarium irreversibly determined during the early postnatal life of the mother, or is each oocyte labile until just before oviposition? In the former case, the probability of an amictic female producing a mictic daughter at any time throughout her reproductive period would reflect the population density she experienced while young and not that at the time she oviposited an egg. 3. Amictic females of two clones of a Florida strain of B. calyciflorus were cultured singly from birth at a low or high density (in a large or small volume) until about halfway through their reproductive period and then switched (experimental treatment), or not (control treatment), to the other density condition. The results indicate that the female fate of an oocyte is determined by maternal population density during oogenesis. Eggs oviposited soon after transfer from low to high density had the same, or a higher, probability of becoming mictic females compared with those produced by control females kept at the high density; eggs oviposited after transfer from the high to the low density had the same low probability of becoming mictic females as those produced by control females kept at the low density. 4. Control females kept at the high density were less likely to produce mictic daughters as they aged. This decline is not because of a decreased propensity of older females to respond to crowding, as older females responded maximally when transferred from a low to a high population density. 5. As oocytes in amictic females respond to maternal population density only during oogenesis, there is a negligible lag between the population‐density signal in the environment and the commitment to sexual reproduction. This minimises the obligatory two‐generation lag between this signal and production of resting eggs, and thus reduces the possibility that crowding will lead to food limitation before production of these eggs.     

孤雌生殖累积世代数和雌体年龄对萼花臂尾轮虫繁殖的影响

采用单个体培养方法,研究了孤雌生殖的累积世代数和雌体年龄对萼花臂尾轮虫混交雌体形成和产卵量的影响,结果表明,随着轮虫孤雌生殖累积世代数和增加,各代中的总混交雌体百分率呈减小的趋势,年轮的雌体可产生较多的混交雌体,非混交雌体所产后代中的总混交雌体百分率具有随其祖母年龄的增大而增大的趋势,孤雌生殖的累积世代数对轮对轮虫非混交雌体的平均产卵量无显著的影响,非混交雌体的年龄对其报代的平均产卵量亦无显著的影响。     

Effect of food concentration on the production and viability of resting eggs of the rotifer Brachionus: implications for the timing of sexual reproduction

1. Life‐table experiments with Brachionus calyciflorus test several hypotheses related to the idea that sexual reproduction in monogonont rotifers should occur when food resources are favourable. 2. The food concentration necessary for a fertilised mictic female to produce one phenotypically normal resting egg was higher than that for an amictic female to produce one daughter. At the lowest concentration of Cryptomonas erosa (1.25 × 10³ cells mL^?1), the lifetime fecundity of these two types of females was 0.9 and 1.4, respectively. 3. The lifetime fecundity of both fertilised mictic females and amictic females increased with food concentration to 3.4 resting eggs and 15.2 daughters female^?1, respectively. The approach to maximal fecundity with increasing food concentration was more rapid for fertilised mictic females, such that their lifetime fecundity relative to that of amictic females gradually decreased from 0.64 (at 1.25 × 10³ C. erosa mL^?1) to 0.22 (at 2.5 × 10⁴ C. erosa mL^?1). 4. The probability of a fertilised mictic female producing one or more abnormal resting eggs during her lifetime was high (approximately 75%). The mean proportion of abnormal eggs produced per female varied among the different food‐concentration treatments (26–38%) but was not higher at the low food concentrations. 5. The proportion of normal resting eggs that hatched was high (51–71%); those produced at low food concentrations were no less likely to hatch than those produced at high food concentrations. No abnormal resting eggs hatched. 6. The probability of a fertilised mictic female producing an abnormal resting egg increased rapidly with her age at all food concentrations. The probability of a normal resting egg hatching declined with maternal age at the low food concentration in one of two experiments. 7. The results support the idea that induction of mictic females should occur when food resources are good. Coincidence of sexual reproduction with low food availability risks low production of resting eggs for several reasons. Population size may be small, with a low probability of encounters between young mictic females and males, and fertilised mictic females may be unable to mature and produce resting eggs.     

Low crowding threshold for induction of sexual reproduction and diapause in a Patagonian rotifer

1. This study investigates the basis and ecological significance of the extremely high propensity for mixis (sexual reproduction) observed in laboratory populations of Brachionus calyciflorus from a temporary pond in Patagonia. 2. Experiments with stem females hatched from resting eggs showed that these females were exclusively amictic but produced mictic daughters even at very low population densities. In six experiments, newly hatched stem females were cultured individually in different volumes (1.5, 12, 40 and 150 mL). The percentage of mictic daughters (mixis ratio) was high in the smaller volumes (e.g. 44–83% in 1.5 mL) and lower in the largest volume (6–21% in 150 mL). A regression analysis combining the data from these experiments showed a highly significant decrease in mixis ratio with volume and indicated that the lowest population density at which mixis still occurs (mixis threshold) would be 3.4 females L^?1 (95% CL 2.9–4.0 females L^?1). This value is considerably lower than mixis thresholds for other rotifers (25–250 females L^?1 for many species and 9000–477 000 females L^?1 for some Spanish Brachionus plicatilis). 3. In three additional experiments, stem females and their amictic daughters were cultured individually in 150 mL. The percentage of mictic daughters produced by these two generations of females was not significantly different, showing that the mixis response to crowding is not inhibited in the stem‐female generation. 4. Laboratory experiments showed that two common predators of the Patagonian B. calyciflorus (the calanoid copepod Parabroteas sarsi and the backswimmer Notonecta vereertbruggheni) each ate 60–70 B. calyciflorus predator^?1 per day and cleared all rotifers from c. 250 mL per day. Thus, a very low mixis threshold and high maximal mixis ratio may ensure production of some resting eggs soon after colonisation of the pond and before complete removal from the plankton. 5. Two laboratory experiments showed that resting eggs of the Patagonian B. calyciflorus hatched at variable rates (28 and 81%) after a brief diapause when kept in the conditions under which they were produced and oviposited (20–21 °C; L: D 16: 8). Early hatching of resting eggs from pond sediment may allow multiple periods of colonisation and resting egg production in a season. This may offset the fitness cost of limited population growth through female parthenogenesis in the face of unpredictable and abrupt risk of extinction because of predators.     

Optimal rates of bisexual reproduction in cyclical parthenogens with density-dependent growth

This work explores theoretical patterns of reproduction that maximize the production of resting eggs and the long-term fitness of genotypes in cyclical parthenogens. Our focus is on density-dependent reproduction as it influences the consequences of a trade-off between producing amictic daughters – which reproduce parthenogenetically and subitaneously – and producing mictic daughters – which undergo meiosis and bisexual reproduction. Amictic females increase competitive ability and allow the population to achieve a larger size; mictic females directly contribute to population survival through harsh periods by producing resting eggs. Although morphologically indistinguishable, the two types of females differ greatly in their ecological and reproductive roles. What factors underlie the differential allocation of resources to produce amictic and mictic females? Using a demographic model based on readily accessible parameters we demonstrate the existence of a frequency of mictic females that will maximize the population's long-term fitness. This frequency, termed the optimal mictic ratio, m_o, is 1 ? (q/b)^1/2, where q is the mortality rate and b is the maximum birth rate. Using computer simulation we compared the fitness of a population with this constant mictic ratio with populations having multiple switches from complete parthenogenetic growth to complete allocation in mixis (mictic ratio either 0 or 1). Two important conclusions for optimal mixis in density-dependent growth conditions are: (1) intermediate mictic ratios are optimal, and (2) optimal mictic ratios are higher when habitat conditions are better. Physiological cues responding to differences in birth and death rates are common so that it is possible that populations may adjust their relative rates of mictic and amictic female production in response to environmentally induced changes to the optimum mictic ratio. Our analysis demonstrates that different patterns of mixis are expected in different type of habitats. Since the optimal mictic ratio is sensitive to the effects of a variety of environmental challenges, our model makes possible a new means to evaluate life history evolution in cyclical parthenogens.     

Environmental and endogenous control of sexuality in a rotifer life cycle: developmental and population biology

Induction of mictic females, and hence initiation of sexuality, in the life cycle of some Brachionus requires an environmental stimulus associated with crowding. The inducing stimulus appears to be a taxonomically specific chemical released into the environment by the rotifers. Oocytes are induced to develop into mictic females before they are oviposited by their amictic mothers and begin cleavage divisions. Thus, the inducer affects the oocyte in the maternal body cavity either directly or indirectly by altering the physiology of its mother. The level of sexual reproduction expressed in populations of a Florida strain of B. calyciflorus is controlled by two types of endogenous factors and by the degree of crowding. First, some fraction of genetically identical oocytes in a clonal population fails to respond to even extreme crowding conditions, thus ensuring some potential for continued population growth by female parthenogenesis. Second, the propensity of amictic females to produce mictic daughters is extremely low when they hatch from fertilized resting eggs and then gradually increases to an asymptote after about 12 parthenogenetic generations. This multigenerational parental effect likely is due to a cytoplasmic factor in fertilized eggs that inhibits expression of the mictic-female phenotype and that is gradually diluted in successive parthenogenetic generations. The effect may increase a clone's genetic contribution to the resting-egg bank by increasing its population size through parthenogenetic generations before mictic females are induced.     

Effect of diazinon on life stages and resting egg hatchability of rotifer Brachionus plicatilis

The effects of organophosphate pesticide, diazinon, on life history parameters and hatchability of resting eggs of rotifer Brachionus plicalitis were assessed. Newly hatched (<1 h-old) neonates were individually cultured in six varying concentrations (0/control, 0.1, 1.0, 2.5, 5.0 and 10.0 mg/l) of diazinon. The life history parameters such as time (h) the rotifers bear first egg and release first neonate, reproductive period, net reproductive rate, mixis, intrinsic rate of population increase, and life span were evaluated. Results showed that among the life history parameters, the time the rotifers took to release neonates is the most sensitive, giving the lowest EC₅₀ value of 1.24 mg/l. The fecundity of maternal females, amictic and mictic daughters was also investigated. Rotifers exposed to 10.0 mg/l produced significantly fewer amictic daughters, and at this concentration, rotifers did not produce any mictic daughter. At 5.0 mg/l, the number of male offspring was significantly lower than the control. Furthermore, the hatchability of resting eggs produced by the rotifers was evaluated when exposed to diazinon: from birth until they produced resting eggs (early development); during late developmental stage of resting eggs (before diapause); and during diapausing stage. The hatchability of the resting eggs was not affected when exposure was timed at late developmental and diapausing stages. Overall results showed that even though amictic females reproduced normally in the presence of low-concentration of diazinon, sexual reproduction is severely affected, especially the hatchability of resting eggs when the exposure was timed on its early developmental stages. Guest editors: S. S. S. Sarma, R. D. Gulati, R. L. Wallace, S. Nandini, H. J. Dumont & R. Rico-Martínez Advances in Rotifer Research     

Sexual reproduction and diapause of Hexarthra sp. (Rotifera) in short-lived ponds in the Chihuahuan Desert

1. In the life cycle of monogonont rotifers it is generally assumed that diapausing eggs invariably hatch into amictic stem females which produce female offspring parthenogenetically. Diapausing eggs are only produced by later generations after sexual reproduction has been induced by environmental cues. 2. We show that populations of an undescribed Hexarthra species inhabiting small temporary ponds in the Chihuahuan Desert deviate from this life cycle pattern. These ponds may dry within days and up to 85% of females were mictic. Females producing male offspring and diapausing eggs were observed 1 or 2 days, respectively, after ponds had filled with water. 3. Under laboratory conditions, 7–46% of females hatching from re‐hydrated sediments were sexual. Male offspring of these females can fertilise other mictic stem females leading to diapausing egg formation. In laboratory experiments, females produced fully developed diapausing eggs within 1.9 days at 20 °C and 1.2 days at 30 °C. 4. In addition, embryonic development time (1.1–0.3 days at temperatures between 12 and 30 °C) and juvenile period (2.1–0.5 days for the same temperature range) are shorter than those of other rotifer species. In short‐lived habitats, the potential for rapid population development and production of new diapausing eggs may be crucial in the long‐term survival of populations.     

Precopulatory mate guarding and mating behaviour in the rotifer Epiphanes senta (Monogononta: Rotifera)

Epiphanes senta is a littoral rotifer species that occurs in temporary waters and displays a mating behaviour which has not, to my knowledge, so far been described for monogonont rotifers. Monogonont rotifers show distinctive periods within their life cycle during which mictic females appear. Mictic females produce haploid eggs that develop into males or into diapausing eggs if fertilized. The females of E. senta are mostly stationary on the substrate while males are more active swimmers. If they encounter eggs with female embryos of their own species, they attend them and mate with the hatching female. Experiments showed that males are able to discriminate between male, female and diapausing eggs. They exhibit a strong preference for female eggs that are only a few hours away from hatching compared with eggs in early developmental stages. Further experiments did not show any significant differences in male attendance of mictic and amictic eggs. It is hypothesized that males judge the age of a female egg by sensing a chemical that is produced by the growing embryo and diffuses through the egg shell. The male mating behaviour is similar to precopulatory mate guarding known from arthropods but it lacks the monopolization of the female by the male.     

Incomplete induction of mixis in <Emphasis Type="Italic">Brachionus calyciflorus</Emphasis>: patterns of reproduction at the individual level

In the rotifer Brachionus calyciflorus mictic-female production is density-dependent and appears to be induced by a chemical—a quorum sensing molecule—produced by the females themselves. Even at the highest densities, however, populations never become entirely mictic: i.e., some amictic females continue to be produced. Surprisingly, the phenomenon also occurs in clonal laboratory populations with genetically identical individuals. Here, we study how this ecologically adaptive phenomenon is generated at the level of individual reproducing females. In a life-history experiment we subjected 123 amictic females of a clone of B. calyciflorus separately to a daily renewed stimulus of culture medium conditioned at a density of 30 females ml⁻¹. For each of these mothers we isolated the lifetime offspring individually and recorded whether these females were amictic or mictic. Mothers produced on average 16 offspring but none of the mothers produced 100% mictic offspring; the average proportion of mictic females was 30%, despite the extremely strong stimulus. The distribution of amictic vs. mictic offspring was not uniform over the mothers’ lifetime. Early and late offspring had a low probability of being mictic whereas mid-aged mothers produced the highest proportion of mictic daughters (up to 56%). We conclude that not all oocytes of B. calyciflorus can be turned into mictic females, even when the mictic-female-inducing stimulus is extremely high. Propensity to become a mictic female also depends on the rank of an egg within a female’s offspring production. Despite these regularities, we observed considerable stochastic variability with respect to individual mothers’ life histories. Guest editors: S. S. S. Sarma, R. D. Gulati, R. L. Wallace, S. Nandini, H. J. Dumont & R. Rico-Martínez. Advances in Rotifer Research.     

Life history characteristics of three types of females in Brachionus calyciflorus Pallas (Rotifera) fed different algae

This study describes the life history characteristics of amictic, unfertilized mictic and fertilized mictic females of the rotifer Brachionus calyciflorus cultured individually on two different algae at 0.1 mg ml^–1 food concentration and 27 °C. The duration of the juvenile period of amictic females was significantly shorter on Chlorella pyrenoidosa Chick than on Scenedesmus obliquus Kütz or both algae together. The duration of the juvenile period of unfertilized mictic females was significantly longer, and the number of eggs produced by amictic females was significantly larger on Chlorella pyrenoidosa than on S. obliquus. When fed the same type of alga, the duration of the juvenile period of the fertilized mictic females was the longest among the three types of females, and the durations of the reproductive period of the amictic females and the post-reproductive period of the fertilized mictic females were longer than, or equal to those of the other two types of females, respectively. The number of eggs produced by an unfertilized mictic female was the largest among the three types of females, and that of amictic females was larger than or equal to that of fertilized mictic females, depending on the type of diet.     

Influence of the age of algae fed to rotifers (Brachionus plicatilis O.F. Müller) on the expression of mixis in their progenies

Summary The sequence of the appearance of mixis in the rotifer Brachionus plicatilis was followed among the descendents of amictic rotifers transferred from a high salinity media (40 S) to a low one (9 S). All the neonates that hatched from the amictic eggs, after being transferred to a low salinity, were amictic. Each one of these neonates was cultured individually and its offspring removed periodically every 8–10 h. It was observed that throughout their reproductive phase, these parental females retained their potential to produce either mictic or amictic offspring. All the first produced neonates developed into amictic females, but among those produced later, three patterns were prevalent. The prevalent pattern (type A) was one in which the probability of a neonate being mictic increased towards the middle of the parents' reproductive phase and was followed by a slow decline. In the second pattern (type B), the probability of a daughter being mictic was constant throughout the parents' reproductive phase. It is suspected that the quality of food supplied to the rotifers determines the appearance of patterns, A, B or C. It is postulated that the innate capacity of rotifers to undergo mixis is genetically controlled, while its expression is modulated by environmental conditions.     

Induction of sexual reproduction and resting egg production in Brachionus Plicatilis reared in sea water

Brachionus plicatilis raised in our laboratory in sea water reproduces asexually even under high crowding conditions (at least 40 individuals per ml). Amictic females were induced to produce mictic females, males and resting eggs by reducing the concentration of the sea water culture medium. Mictic females and males appeared predominantly among the progeny produced by the amictic females during 4 days following their transfer into 25% sea water. Resting eggs appeared first 5–12 days after the onset of the experiment. Following the disappearance of males, the culture consisted of amictic females.Resting eggs produced by the method described above may be preserved for at least three months at –14°C or by desiccation at room temperature. Under the appropriate experimental conditions, resting eggs hatch into amictic females. Since B. plicatilis is one of the most commonly used food sources of fish larvae in aquaculture, the methods reported here may offer an easy and versatile way of preserving rotifer culture stock to be used on demand.     

The significance of mating processes for the genetics and for the formation of resting eggs in monogonont rotifers

Crossbreeding experiments with three geographically distinct strains (E, S, and L) of the rotifer Brachionus plicatilis have been carried out in an attempt to elucidate the apparent male sterility of strain L, which is unable to produce resting eggs.The 9 crossing possibilities of the three strains have been investigated in 27 experiments. The results refute the concept of male sterility. L-males copulate successfully with mictic as well as with amictic females of strains E and S. Fertilized amictic E and S females produce defective resting eggs, which have only one thin shell and, which disintegrate after deposition. L-females cannot, as a rule, be fertilized. Moreover, crosses between strain E and S are succesful only in one direction; the reciprocal crosses failed.To explain the present results a hypothesis is suggested that the thickness of the body wall of newborn females differs in the three strains, and between mictic and amictic individuals. A sequence of gradually increasing body wall thickness of all types of females involved, together with a comparably increasing penetration ability of the males of strain E, S, and L explains the success or failure of all crosses, including the unilateral cross E × S.     

Production of the rotifer Brachionus plicatilis (Ploimida: Brachionidae) in a Brazilian coastal lagoon

Secondary production, density and biomass fluctuations, and some reproductive aspects of the rotifer Brachionus plicatilis were studied weekly for two years in a Brazilian coastal lagoon. Food (and secondarily temperature) seemed to be the main factor involved in the summer increments. In the first summer, a mass mortality of fish caused changes in the phytoplankton, which switched from the dominance of Cyanophyceae to Chlorophyceae and other groups. For two months, production, density and biomass were relatively high until the recovery of the Cyanophyceae dominance. In the second summer, peaks of shorter duration and higher values of production, in comparison to the first one, were found. The contribution of resting eggs hatching, in the first summer, and the increment in parthenogenetic reproduction, in the second summer, would account for the population increases observed. Males and mictic females occurred when densities were high.     

A study of the freshwater rotifer Brachionus calyciflorus in Pekanbaru, Riau, Indonesia

Brachionus calyciflorus is one of freshwaterrotifers found in fish ponds in Pekanbaru, Riau,Indonesia. Its density varied depending on that ofphytoplankton. Maximum of 2.5 ×10³ ind. l^–1 was found in ponds fertilised with animal wastes.The fecundity and population growth of B.calyciflorus was studied. Results indicated that the fecundity of amictic females was higher thanthat of unfertilized mictic females. During their life span, amictic and mictic females produced29.7 and 12.5 eggs per female, respectively. In the mass culture of 500 ml media, the highestdensity of female and male rotifers was 975.8 and 9.6 ind. ml^–1, respectively. During 8 days cultureperiod, they also produced eggs as many as 124.2 ml^–1.Human and animal wastespromoted the growth of phytoplankton as food forrotifers in the pond. A laboratory study confirms this. The best growth of B. calyciflorus at a density of 109 ind. ml^–1 was found at 0.5 g l^–1 of humanexcreta. A high density of B. calyciflorus (542ind. ml^–1) was also found in semi-continuous culturewith chicken excreta.     

Delayed mixis in rotifers: an adaptive response to the effects of density-dependent sex on population growth

In most cyclically parthenogenetic life cycles, sex is neededto produce resting stages. In several species of cyclicallyparthenogenetic rotifers, some generations of clones are notresponsive to a density-dependent signal that triggers sexualfemale production. These unresponsive rotifers hatch from restingeggs and typically pass 8–12 generations of female parthenogenesisbefore becoming receptive to the mixis signal. We addressedthe selection for mixis delay using a simulation model. A delayof sexual reproduction could increase population growth throughparthenogenesis and thus the number of resting eggs ultimatelyproduced. In a monomorphic population without mixis delay, wedetermined the optimal ratio of mictic to amictic females (mixisratio) to be 45%, and the optimal population density thresholdfor induction of mictic females (mixis threshold) to be 82 rotifersL^–1. This mixis pattern, however, was not an evolutionarilystable strategy. A mixis ratio of 14% and threshold of 70 rotifersL^–1 proved to be resistant to invasion by other mixispatterns. When we gave this phenotype a mixis delay of 8–10days, it could invade a population with the same mixis pattern,but lacking a mixis delay. The advantage of delaying mixis wasrelatively small, suggesting that a polymorphism is possible.     

Dietary tocopherol and sexual reproduction in the rotifers Brachionus calyciflorus and Asplanchna sieboldi.

Brachionus calyciflorus contained no detectable tocopherol (vitamin E) when cultured for long periods on the yeast Rhodotorula glutinis. Using a spectrofluorometric assay and the Asplanchna body-wall-outgrowth response bioassay the limits of detection were 6.4 ng and 3.78 pg alpha-tocopherol per g dry weight Brachionus, respectively. These tocopherol-free Brachionus reproduced normally both parthenogenetically and sexually, producing active and potent males and fertilized resting eggs which developed into viable young rotifers. The apparent lack of a tocopherol requirement for male fertility in B. calyciflorus probably also applies to Asplanchna sieboldi and A. brightwell -- rotifers whose male-producing (mictic) females are induced by dietary tocopherol and whose males therefore are never tocopherol-deficient. The adaptive significance of the tocopherol requirement for the initiation of sexual reproduction in Asplanchna sieboldi--formerly hypothesized to be the result of a tocopherol requirement for male fertility--is now thought to be related to the role of tocopherol in controlling female polymorphism in this species.     

Effect of incubation and preservation on resting egg hatching and mixis in the derived clones of the rotifer Brachionus plicatilis

The marine rotifer Brachionus plicatilis typicus (Clone 8105A, Univ. of Tokyo) was cultured in 500 ml beakers to form resting eggs. Tetraselmis tetrathele was used as a culture food. Just after formation, resting eggs were exposed to various temperature (5–25 °C) and light regimes (24L: OD and OL : 24D). When eggs were exposed to light just after formation, the eggs hatched sporadically over a month. No hatching was observed for six months when eggs were preserved under dark conditions regardless of the temperature. These eggs hatched simultaneously after being exposed to light and eggs preserved at 5 °C showed twice as high hatching rate (40%) as that of eggs preserved at 15–25 °C (24%). Clones from resting eggs that were kept under different temperature and light regimes were reared individually to the third generation. Incubation at 25 °C with lighting produced the highest (5.4% and 5.2 %) rate of mictic females during their 2nd and 3rd generations, respectively. The lowest rates (0 and 1.5%) were found when the eggs were kept at 5 °C in total darkness for six months. A lower rate of amictic female production was found in clones with higher rates of mixis.