Germ-soma differentiation in volvox.

Volvox carteri is a spherical green alga with a predominantly asexual mode of reproduction and a complete germ-soma division of labor. Its somatic cells are specialized for motility, incapable of dividing, and programmed to die when only a few days old, whereas its gonidia (asexual reproductive cells) are nonmotile, specialized for growth and reproduction, and potentially immortal. When a gonidium is less than 2 days old it divides to produce a juvenile spheroid containing all of the somatic cells and gonidia that will be present in an adult of the next generation. The first visible step in germ-soma differentiation is a set of asymmetric cleavage divisions in the embryo that set apart small somatic initials from their large gonidial-initial sister cells. Three types of genes have been found to play key roles in germ-soma specification. First a set of gls genes act in the embryos to shift cell-division planes, resulting in the asymmetric divisions that set apart the large-small sister-cell pairs. Then a set of lag genes act in the large cells to prevent somatic differentiation, while the regA gene acts in the small cells to prevent reproductive development. An inducible transposon was used to tag and recover some of these and other developmentally important genes. The glsA gene encodes a chaperone-like protein that, like another chaperone that is one of its putative binding partners, is associated with the cell division apparatus, although how this leads to asymmetric division remains to be elucidated. The regA gene encodes a somatic-cell-specific nuclear protein that appears to function by repressing genes required for chloroplast biogenesis, thereby preventing somatic cells from growing enough to reproduce. Somatic-cell-specific expression of regA is controlled by three intronic enhancers.     

The evolutionary ecology (evo-eco) of plant asexual reproduction

The different modes of asexual plant reproduction are reviewed in the context of the evolutionary ecology (evo-eco) of retaining old (and colonizing new) habitats and exploring new ones. The modes of asexual reproduction are contrasted between the green algae and the nonvascular and vascular land plants (bryophytes and tracheophytes, respectively). The chronology of their appearance is also traced in the fossil record. Among bryophytes, the indeterminate growth of protonema and the subsequent fragmentation of gametophytes provide a successful asexual reproductive stratagem, one that is paralleled by indeterminate rhizomaty and subsequent fragmentation of tracheophyte sporophytes. Self-compatible (autogamous) bisexual (monoicous) gametophytes capable of producing viable homozygous sporophytes offer a method for colonizing new sites by means of a single spore, a stratagem that is paralleled in tracheophyte sporophytes by the co-option of sexual reproductive structures (e.g., flowers) to produce diploid embryos asexually (e.g., variants of apomixis such as nucellar embryony). A review of the fossil record and the biology of extant taxa show that some of these modes of asexual reproduction are ancient and present in the green ancestors of the land plants (i.e., the charophycean algae). An overview of the data indicates that asexual reproduction provides a bet-hedging stratagem that can obviate some of the many costs of sexual reproduction, and preserve well-adapted genomes within previously occupied sites.     

Interference of asexual and sexual reproduction in the green hydra

The green hydra, Hydra viridissima, has three sexes: hermaphrodite, male, and female. I investigated the reproductive strategies of the green hydra and the relationship between asexual budding and sexual reproduction. The proportion of mature individuals in the asexually reproducing population increased with increasing temperature. Sexual reproduction did not interrupt asexual budding in hermaphrodites or males; sexual reproduction did interrupt asexual budding in females. Sexual reproduction also resulted in exponential population growth during the reproductive season. The number of asexual buds on each parental individual was positively correlated with the parental individual size in asexual individuals and in males. The same positive correlation was found between the number of testicles and the size of males. These correlations reflect a common tendency in asexual and sexual reproduction: larger parental individuals have a greater number of propagules or gametes. No correlation was found between size and buds or size and gonads in hermaphrodites; hermaphrodites had at most one asexual bud and were significantly larger than males, females, and asexual individuals. The larger size of hermaphrodites supports the hypothesis that producing both female and male gonads is more energetically costly than producing only one type of gamete (gonochorism).     

Food quality triggers the reproductive mode in the cyclical parthenogen <Emphasis Type="Italic">Daphnia</Emphasis> (Cladocera)

Cyclic parthenogenesis (heterogony) is a widespread reproductive mode found in diverse taxa such as digenean trematodes, gall wasps, gall midges, aphids, cladocerans and rotifers. It is of particular interest as it combines the advantages of asexual reproduction (rapid population growth) and sexual reproduction (recombination). Usually sexual reproduction is initiated when, or slightly before, environmental conditions deteriorate, and often results in the production of resting stages. The optimal timing of diapause induction must thus be under strong natural selection. Using the cladoceran Daphnia as a model system, we show here for the first time that the switch from parthenogenetic to sexual reproduction in a cyclical parthenogenetic organism can be influenced by the chemical composition of food. Under crowding conditions Daphnia reproduced parthenogenetically with subitaneous eggs when fed the algal species Cryptomonas sp., but started the production of resting eggs when fed with the green algal species Scenedesmus obliquus. Supplementation experiments with lipids and especially proteins showed that the induction of resting egg production in two clones of different Daphnia species was due to a dietary deficiency in the green alga. Hence, the low food quality induced a switch in the reproductive mode that may contribute to optimal timing of the sexual reproduction of Daphnia in nature. Furthermore, our results have two other major implications: first, they suggest that protein compounds should be added to the list of diet constituents potentially limiting or influencing Daphnia reproduction. Second, we show that the role of food quality goes far beyond the up to now documented effects of food quality on somatic growth and trophic transfer efficiency of herbivores: due to its effects on sexual reproduction and the production of resting eggs, food quality might influence genetic diversity and long-term persistence of Daphnia in lakes. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The eco-evolutionary importance of reproductive system variation in the macroalgae: Freshwater reds as a case study

The relative frequency of sexual versus asexual reproduction governs the distribution of genetic diversity within and among populations. Most studies on the consequences of reproductive variation focus on the mating system (i.e., selfing vs. outcrossing) of diploid-dominant taxa (e.g., angiosperms), often ignoring asexual reproduction. Although reproductive systems are hypothesized to be correlated with life-cycle types, variation in the relative rates of sexual and asexual reproduction remains poorly characterized across eukaryotes. This is particularly true among the three major lineages of macroalgae (green, brown, and red). The Rhodophyta are particularly interesting, as many taxa have complex haploid–diploid life cycles that influence genetic structure. Though most marine reds have separate sexes, we show that freshwater red macroalgae exhibit patterns of switching between monoicy and dioicy in sister taxa that rival those recently shown in brown macroalgae and in angiosperms. We advocate for the investigation of reproductive system evolution using freshwater reds, as this will expand the life-cycle types for which these data exist, enabling comparative analyses broadly across eukaryotes. Unlike their marine cousins, species in the Batrachospermales have macroscopic gametophytes attached to filamentous, often microscopic sporophytes. While asexual reproduction through monospores may occur in all freshwater reds, the Compsopogonales are thought to be exclusively asexual. Understanding the evolutionary consequences of selfing and asexual reproduction will aid in our understanding of the evolutionary ecology of all algae and of eukaryotic evolution generally.     

Asexual reproduction, habitat colonization and habitat maintenance in bryophytes

Asexual reproduction s. l. is widespread in plants and also a basic reproductive mechanism in bryophytes. Today, three types of asexual reproduction are recognized: (1) the asexual reproduction s. str. by regeneration from ± specialized caducous organs (leaves, leaf apices, shoots, branches, bulbils) and by production of specialized propagules (gemmae, protonemal brood cells, tubers), (2) fragmentation of plants, resp. part of plants into ± unspecialized fragments, and (3) clonal reproduction (cloning). The latter occurs in bryophytes by protonema decay, by disintegration of modules, resp. formation of ramets (dividuals, “daughter plants”) that leads to self-cloning or forced-cloning of parts of the gametophyte (shoots, stoloniferous and rhizomatous axes, rhizoid wicks, basiscopic innovation plants). Clonal reproduction (cloning), in former time scarcely noted, gained great interest within the last decade mainly in vascular plants showing clonal growth. This reproduction mechanism is thought to be a keystone factor for asexual reproduction, habitat colonization and habitat maintenance. Species which reproduce clonally are able to colonize and maintain habitats in an effective way by the so-called “consequent vegetative multiplication”. The review presents an overview of the current state of knowledge of asexual reproduction types in bryophytes, with a focus on fragmentation and clonal reproduction (cloning), the mechanisms of habitat colonization and habitat maintenance, which all are of important significance in the dynamic processes of development of bryophyte populations.     

A New Species of Stigeoclonium (Chaetophoraceae,Ulotrichales)

This paper describes a new species, Stigeoclonium ovisporum Wei (Chaetophoraceae, Ulotrichales), from Yunnan and Sichuan provinces, China. In asexual reproduction of this alga, narrow oviod sporangia produce zoospores. The differentiation of vegetative cellsand reproductive ones is an advanced character in the genrus Stigeoclonium.     

毛枝藻属一新种

本文报道了丝藻目、胶毛藻科、毛枝藻属的一新种—卵孢毛枝藻,此种采自我国云南省和四川 省。此藻类由狭卵形的孢子囊产生动孢子进行无性生殖,在毛枝藻属的生殖中,营养细胞和生殖细胞分化是一种进化类型。     

Variations in sexual and asexual reproduction of Scirpus mariqueter along an elevational gradient

In order to assess the importance of sexual and asexual reproduction during the life history of Scirpus mariqueter, its reproductive and growth characters were concurrently examined along an elevational gradient (from low elevation to high elevation). The proportions of flowering shoot and inflorescence mass, seed : flower ratio and seed weight were used to quantify the investment in sexual reproduction. The proportions of current-year shoot and rhizome mass were used to quantify the investment in asexual reproduction, and the proportion of corm mass was used for growth, respectively. It was found that vegetative propagation predominated at low elevation, whereas sexual reproduction predominated at high elevation; and that sexual reproduction increased with declining asexual reproduction along the gradient. The results suggest that asexual reproduction is relatively favored in the early life stage, whereas sexual reproduction is favored when the population becomes mature and aged, probably because of the functional differentiation between the two reproductive types. Sexual productive characters (i.e. the proportions of flowering shoot and inflorescence mass) were negatively correlated to both growth and asexual reproductive characters along the gradient, indicating there might exist some trade-offs among growth, sexual and asexual reproduction during the life history. However, no obvious pattern was found between asexual reproductive characters and growth characters along the elevational gradient, possibly because of the varied relationships between them at different life stages. The variations in sexual and asexual reproduction in the species and the relationship between them are thought to be of great significance for local population growth, species persistence and evolution.     

Food quality controls reproduction of the zebra mussel (Dreissena polymorpha)

Species such as Dreissena polymorpha sometimes contribute substantially in the transfer of primary to secondary production. During the ontogenetic cycle, the reproductive investment of adult mussels is one of the main parameters that affect recruitment success. We studied how food quality and temperature affect the reproductive investment in term of egg mass of D. polymorpha in a lake by sampling mussels monthly from 4 m and 15 m depths. Temperature affected reproduction directly and also indirectly through the food. To assess whether temperature and/or food conditions led to the differences observed in mussels sampled from the two depths, mussels were reared in the laboratory under two different temperature regimes for 3 months, simulating the temperature of the lake at 4 m and 15 m depth. Possible effects of food quality were tested at each temperature using four diets differing in fatty acid composition. Temperature played an important role as a trigger for spawning, and the type of diet clearly affected the reproductive investment. When the heterokont chromophyte alga Nannochloropsis limnetica, which is rich in polyunsaturated fatty acids (PUFAs) and long-chained PUFAs (>C18), was fed to mussels, an increased egg mass was obtained. This result was in contrast to that found when the green alga Scenedesmus obliquus and the cyanobacterium Aphanothece sp., both of which are deficient in long-chained PUFAs, were offered as food to the mussels. Such a PUFA-dependent food quality may affect reproduction in lakes. Food quality effects vary seasonally in a lake and may be most important in summer, when low-food-quality green algae and cyanobacteria are abundant. The low biochemical quality of these blooms may affect at least the later period of gametogenesis of D. polymorpha, which reproduces from June to August.     

Parthenogenetic female populations in the brown alga Scytosiphon lomentaria (Scytosiphonaceae,Ectocarpales): decay of a sexual trait and acquisition of asexual traits

In isogamous brown algae, the sexuality of populations needs to be tested by laboratory crossing experiments, as the sexes of gametophytes are morphologically indistinguishable. In some cases, gamete fusion is not observed and the precise reproductive mode of the populations is unknown. In the isogamous brown alga Scytosiphon lomentaria in Japan, both asexual (gamete fusion is unobservable) and sexual populations (gamete fusion is observable) have been reported. In order to elucidate the reproductive mode of asexual populations in this species, we used PCR‐based sex markers to investigate the sex ratio of three asexual and two sexual field populations. The markers indicated that the asexual populations consisted only of female individuals, whereas sexual populations are composed of both males and females. In culture, female gametes of most strains from asexual populations were able to fuse with male gametes; however, they had little to no detectable sexual pheromones, significantly larger cell sizes, and more rapid parthenogenetic development compared to female/male gametes from sexual populations. Investigations of sporophytic stages in the field indicated that alternation of gametophytic and parthenosporophytic stages occur in an asexual population. These results indicate that the S. lomentaria asexual populations are female populations that lack sexual reproduction and reproduce parthenogenetically. It is likely that females in the asexual populations have reduced a sexual trait (pheromone production) and have acquired asexual traits (larger gamete sizes and rapid parthenogenetic development).     

不同生境中克隆植物的繁殖倾向

对克隆植物在林下、草原、荒漠、冻原、农田和水域等6种不同类型生境中的繁殖对策及其生态适应意义进行了归纳和总结。对克隆植物有性繁殖与无性繁殖之间的权衡关系及相对重要性随水分、光照和温度的变化规律进行了阐述。分析表明,在环境条件严酷地区,克隆植物的无性繁殖在种群更新中的相对重要性有所增加。对两种繁殖方式的生态功能特性进行比较,综述了有关二者权衡关系的研究成果及影响二者权衡关系的部分因素。从研究手段上对克隆植物繁殖对策的研究进行了思考与展望。     

Twenty-seven modes of reproduction in the obligate lichen symbiosis

Fungi exhibit some of the greatest reproductive diversity across Eukaryotes. In addition to sexual and asexual reproduction, fungi engage in parasexual (mitotic recombinatorial) processes to acquire new genetic variation. Reproduction has been studied extensively in numerous free-living fungi but comparatively less knowledge exists for lichenized fungi, which are assumed to reproduce only through sexual spores, asexual conidia, and specialized asexual lichen propagules. We present a new conceptual framework describing reproductive modes in lichens that includes sexual and asexual processes as well as accommodating the possibility of parasexual reproduction. To support the plausibility of some of these modes of reproduction, we reviewed data spanning more than 200 years of anatomical investigation. We recovered evidence supporting the possibility of 22 of 27 possible modes of reproduction and found no counter-evidence to suggest the remaining five do not occur in nature. This conceptual framework allows for a greater plurality of reproductive processes than previously acknowledged in lichens, exceeding that of their non-lichenized relatives.     

Effect of temperature and irradiance on the growth and reproduction of the green macroalga, Chaetomorpha valida (Cladophoraceae, Chlorophyta)

“Green tides” are vast accumulations of unattached green macroalgae associated with eutrophicated marine environments. It had major ecological and economic impacts globally, thus understanding of their detailed conditions was required to inform management decisions. Chaetomorpha valida (Cladophoraceae, Chlorophyta), as a fouling green alga, was found in aquaculture ponds along the coast of Dalian and Rongcheng cities, People’s Republic of China in the past few years. Seasonal abundances of this macroalga in China have presented a significant nuisance and caused great loss to the local aquaculture. Using unialgal cultures, the effect of temperature and irradiance on growth and reproduction were investigated. Healthy growth and reproduction occurred in the range of 17–29°C while the upper lethal limit was at 33°C. A suitable temperature range over 21–29°C and a relatively high irradiance of 108?μmol photons?m^?2?s^?1 were more favorable for growth and reproduction. According to our observations, the diversity of reproductive approaches and wide adaption to temperature can be considered to be key factors that facilitate its excessive growth and colonization in spring and summer. Our results provide a basis for establishing a forecast system and taking remediation measures against seasonal green tides of this macroalga.     

Preliminary crystallographic data for plastocyanins from an alga (Enteromorpha prolifera) and from cucumber (Cucumis sativus)

The plastocyanins from a green alga (Enteromorpha prolifera) and cucumber (Cucumis sativus) have been crystallized. Crystal data are as follows: E. prolifera plastocyanin, space group I4, a = b = 53.9 A, c = 59.4 A, Z = 8; C. sativus plastocyanin, space group P4(1) (or P4(3) ), a = b = 66.7 A, c = 46.0 A, Z = 8. Accordingly, the asymmetric units of the crystals contain one and two molecules, respectively.     

Admixed sexual and facultatively asexual aphid lineages at mating sites

Cyclically parthenogenetic organisms may have facultative asexual counterparts. Such organisms, including aphids, are therefore interesting models for the study of ecological and genetic interactions between lineages differing in reproductive mode. Earlier studies on aphids have revealed major differences in the genetic outcomes of populations that are possibly resulting mostly either from sexual or from asexual reproduction. Besides, notable gene flow between sexual and asexual derivatives has been suspected, which could lead to the emergence of new asexual lineages. The present study examines the interplay between these lineages and is based on analyses of population structure of individuals that may contribute to the pool of sexual reproductive forms in the host alternating aphid Rhopalosiphum padi. Using a Bayesian assignment method, we first show that the sexual forms of R. padi on mating sites encompass two genetically distinct clusters of individuals in the western part of France. The first cluster included unique genotypes of sexual lineages, while the second cluster included facultatively asexual lineages in numerous copies, the reproductive mode of the two clusters being confirmed by reference clones. Sexual reproductive forms produced by sexual and facultatively asexual lineages are thus admixed at mating sites which gives a large opportunity for the two clusters to mate with each other. Nevertheless, this study also highlights, as previously demonstrated, that the two clusters retained high genetic differentiation. Possible explanations for the inferred limited genetic exchanges are advanced in the discussion, but further dedicated investigations are required to solve this paradox.     

光周期对中国水螅种群增长、无性出芽生殖及抗氧化酶活力的影响

单细胞真核绿藻在中国水螅(Hydra sinensis)内胚层皮肌细胞中共生是有较高科研价值的特殊生物学现象。水螅宿主细胞为共生藻提供CO2、氮源及矿物质,而共生藻通过光合作用可能为宿主提供碳水化合物等有机物营养,因此水螅与共生藻间代谢流是以共生藻光合作用为中心,但基于代谢流二者间的互作机制目前尚未阐明。水螅通过营养积累进行出芽生殖,从母体脱落的芽体数量间接反映水螅营养积累的相对量。而光暴露时长能影响共生绿藻光合作用,如果共生藻的确能向水螅细胞转移光合作用产物,那光暴露时长应该能间接影响水螅的营养积累、从而进一步影响水螅无性出芽生殖。为证实该假说,本研究应用种群累积培养法,观察了光周期对中国水螅种群增长、无性出芽生殖及抗氧化酶(SOD和CAT)活力的影响。结果显示,光周期对中国水螅种群增长具有明显的影响。培养15 d后,所有实验组水螅的种群密度均为正增长,其中8L∶16D(在一个24h周期内光暴露8 h、黑暗16 h)实验组种群密度最大、而0L∶24D(持续黑暗)实验组种群密度最小。另外,随着光暴露时长的增加,中国水螅SOD及CAT活力整体均呈下降趋势。结果表明,从光周期对中国水螅无性出芽生殖及两种抗氧化酶活力的影响来看,中国水螅对光周期的生理学响应较为敏感,这个现象可能源于共生绿藻能通过向宿主细胞转移光合作用产物的方式为水螅提供营养补充。     

Could different environmental conditions select for different flight characteristics in sexual and asexual parasitoid Venturia canescens?

In the parasitoid Venturia canescens Gravenhorst (Hymenoptera: Ichneumonidae), asexual and sexual wasps coexist in the field in the Mediterranean basin, but only the asexual strain is present indoors. The sexual strain dominates outdoors despite the demographic costs associated with the production of males and mate location. The present study tests whether females of the sexual and asexual strains of V. canescens differ in flight characteristics in line with the differences of their preferred habitats and enquires whether these differences might contribute to the persistence of sexually reproducing individuals in the vicinity of asexual counterparts. The results show that sexual female wasps are smaller than their asexual counterparts. The size of wasps has a strong influence on flight parameters, with larger animals generally being better fliers. In wasps of approximately the same size, sexual wasps fly faster than their asexual counterparts under experimental laboratory conditions, in terms of both the average speed over the observation period as well as the longest single flight. Sexual wasps also perform fewer flights to cover the same distance. Sexual wasps have higher wing loading than asexual ones of the same size, which could have contributed to the observed differences in speed between individuals of both reproductive modes. There are no significant differences between the two reproductive modes in the parameters related to the distance traversed or the time spent in flight. This study shows clear differences in the flight behaviour of sexual and asexual V. canescens. Together with previous results, this finding suggests differential adaptations to their preferred habitats. These differences might ease the competition between modes of reproduction through niche and habitat differentiation and might help to explain their coexistence on a geographical scale.     

The effect of temperature on infradian rhythms of reproduction in <Emphasis Type="Italic">Ulva fenestrata</Emphasis> postels et ruprecht, 1840 (Chlorophyta: <Emphasis Type="Italic">Ulvales</Emphasis>)

The influence of temperature on the duration of infradian (over 28 h) rhythms of reproduction in the green alga Ulva fenestrata was studied in the laboratory under controlled conditions combining four temperatures (5, 10, 15, and 20°C), two irradiance levels (40 and 60 μE/(m² s)) and neutral daylengths (12: 12 h light: dark). The rise in water temperature explained 78.4% of the reduction in the reproduction cycles (from 30 to 5 days). It is suggested that U. fenestrata has an endogenous reproductive rhythm with a period of 5 days. In unfavorable conditions, one or more reproduction cycles are omitted, and formation of gametes or spores attains a periodicity of 10, 15, or more days.     

Culture and hybridization experiments on an ulva clade including the Qingdao strain blooming in the yellow sea

In the summer of 2008, immediately prior to the Beijing Olympics, a massive green tide of the genus Ulva covered the Qingdao coast of the Yellow Sea in China. Based on molecular analyses using the nuclear encoded rDNA internal transcribed spacer (ITS) region, the Qingdao strains dominating the green tide were reported to be included in a single phylogenetic clade, currently regarded as a single species. On the other hand, our detailed phylogenetic analyses of the clade, using a higher resolution DNA marker, suggested that two genetically separate entities could be included within the clade. However, speciation within the Ulva clade has not yet been examined. We examined the occurrence of an intricate speciation within the clade, including the Qingdao strains, via combined studies of culture, hybridization and phylogenetic analysis. The two entities separated by our phylogenetic analyses of the clade were simply distinguished as U. linza and U. prolifera morphologically by the absence or presence of branches in cultured thalli. The inclusion of sexual strains and several asexual strains were found in each taxon. Hybridizations among the sexual strains also supported the separation by a partial gamete incompatibility. The sexually reproducing Qingdao strains crossed with U. prolifera without any reproductive boundary, but a complete reproductive isolation to U. linza occurred by gamete incompatibility. The results demonstrate that the U. prolifera group includes two types of sexual strains distinguishable by crossing affinity to U. linza. Species identification within the Ulva clade requires high resolution DNA markers and/or hybridization experiments and is not possible by reliance on the ITS markers alone.