THE LIFE CYCLE OE SARGASSUM HORNERI (PHAEOPHYTA) IN LABORATORY CULTURE1

The life cycle of the large dioecious alga Sargassum horneri (Turner) C. Agardh was completed in unialgal culture by controlling photoperiod in relation to the phase of growth. Embryos isolated from a naturally grown female thallus gave rise to early germlings that rapidly formed blades under both short-day (9 h L) and long-day (15 h L) conditions at 20° C Shoot elongation, which followed early blade formation, occurred under the short-day conditions hut not under the long-day conditions. Functional female and male receptacles developed when thalli 8–14 cm long grown under the short-day conditions were transferred to the long-day conditions; gamete fusion occurred when male and female thalli were grown together. Fertilized oospores gave rise to normal thalli in a manner similar to that for in situ plants. Thus, the life cycle of S. horneri was completed in laboratory culture.     

Phenology of a dioecious population of Batrachospermum delicatulum (Batrachospermales, Rhodophyta) in a stream from southeastern Brazil

The seasonal dynamics of a dioecious population of Batrachospermum delicatulum (Skuja) Necchi and Entwisle was evaluated biweekly during the growth period of the macroscopic gametophyte, from late autumn (May) to early spring (October) in a third-order stream from the northeast region of Sao Paulo State, southeastern Brazil (20?43'S, 49?13'W). The population fluctuated throughout the study period in terms of percentage cover, frequency and chlorophyll content. Percent cover and frequency showed a clear pattern with the lowest values at the initial and final stages of the growth period and the highest in July-August (winter). Gametophyte growth was associated with high illumination and low temperature, which agrees with most observations on Batrachospermales in stream environments. Relatively wide variations in reproductive characteristics were observed during the growth period, which were not correlated with percentage cover, frequency and chlorophyll content. The phenologic pattern observed in this population was characterized by a synchronic development of female/male plant ratio with the number of spermatangia per plant resulting in: (i) a higher proportion of fertilized (carpo-sporophytic) plants associated with more male plants; (ii) higher fertilization rates during the periods with higher production of spermatangia per plant; and (iii) higher production of spermatangia per plant when the population had more male plants. These characteristics were largely associated with water temperature and ion content. In terms of reproductive success, the population studied can be regarded as highly efficient, considering the relatively low proportion of male to female plants and high fertilization rates. These data are consistent with a hypothesis to explain high fertilization rates in dioecious species in lotic habitats consisting of outcrossing among adjacent plants with intermingled male and female branches within an algal cluster. Efficient reproductive strategies have been reported in dioecious, monoecious and even mixed populations of B. delicatulum and can be interpreted as adaptations to successfully colonize streams with different characteristics.     

A molecular and morphological investigation of Batrachospermum arcuatum (Batrachospermales, Rhodophyta) in China

Batrachospermum arcuatum specimens were analysed from seven stream segments in North China. Morphological characteristics were observed and cluster analysis was used to evaluate the divergence among thalli from. Sequence data of the rbcL gene (chloroplast gene) and cox2-3 spacer region (mitochondrial gene) were also utilized to evaluate genetic variation in specimens among stream segments. The specimens from four of the streams were monoecious, while the individuals at the other three locations were dioecious. Cluster analysis showed that the monoecious specimens were not separated from the dioecious specimens, based on morphology, but rather the specimens were grouped by geographical closeness and habitat similarity. Likewise, the combined analyses of rbcL and the cox2-3 spacer data from provided more evidence that breeding system (monoecy vs. dioecy) is not a good morphological character to distinguish species.     

Sex ratio from germination through maturity and its reproductive consequences in the liverwort Sphaerocarpos texanus

Summary The dioecious winter ephemeral liverwort, Sphaerocarpos texanus disperses spore tetrads consisting of two males and two females. I examined the subsequent sex ratio of S. texanus at different stages in its life cycle to detect possible mechanisms affecting deviations from a 1:1 sex ratio and the effect of sex ratio on reproductive success. As S. texanus occurs in pure male, pure female and mixed sex clumps, I examined the proportions and sizes of these, the reproductive success of pure female and mixed sex clumps in the field and the sex ratio of germinating plants in a growth chamber. In both the field and the growth chamber the most abundant clump type was pure female followed by mixed sex and pure male clumps. These abundance patterns suggest that males have a lower survival rate than females before emergence and this lower survival rate continues through the gametophytic stage. This disadvantage may be due to the higher susceptibility of males to environmental conditions, to their competitive inferiority to females, and/or to differential resource allocation to the sexes within the spore tetrad. The female biased sex ratio at germination is consistent with predictions from sex ratio theory. Further my field data indicate that males may gain a survival benefit from growing in a mixed sex clump and both males and females benefit reproductively when they occur in mixed sex clumps.     

Clonal genetic structure and diversity in populations of an aquatic plant with combined vs. separate sexes

Clonality is often implicated in models of the evolution of dioecy, but few studies have explicitly compared clonal structure between plant sexual systems, or between the sexes in dioecious populations. Here, we exploit the occurrence of monoecy and dioecy in clonal Sagittaria latifola (Alismataceae) to evaluate two main hypotheses: (i) clone sizes are smaller in monoecious than dioecious populations, because of constraints imposed on clone size by costs associated with geitonogamy; (ii) in dioecious populations, male clones are larger and flower more often than female clones because of sex‐differential reproductive costs. Differences in clone size and flowering could result in discordance between ramet‐ and genet‐based sex ratios. We used spatially explicit sampling to address these hypotheses in 10 monoecious and 11 dioecious populations of S. latifolia at the northern range limit in Eastern North America. In contrast to our predictions, monoecious clones were significantly larger than dioecious clones, probably due to their higher rates of vegetative growth and corm production, and in dioecious populations, there was no difference in clone size between females and males; ramet‐ and genet‐based sex ratios were therefore highly correlated. Genotypic diversity declined with latitude for both sexual systems, but monoecious populations exhibited lower genotypic richness. Differences in life history between the sexual systems of S. latifolia appear to be the most important determinants of clonal structure and diversity.     

Slow evolution of sex‐biased genes in the reproductive tissue of the dioecious plant Salix viminalis

The relative rate of evolution for sex‐biased genes has often been used as a measure of the strength of sex‐specific selection. In contrast to studies in a wide variety of animals, far less is known about the molecular evolution of sex‐biased genes in plants, particularly in dioecious angiosperms. Here, we investigate the gene expression patterns and evolution of sex‐biased genes in the dioecious plant Salix viminalis. We observe lower rates of sequence evolution for male‐biased genes expressed in the reproductive tissue compared to unbiased and female‐biased genes. These results could be partially explained by the lower codon usage bias for male‐biased genes leading to elevated rates of synonymous substitutions compared to unbiased genes. However, the stronger haploid selection in the reproductive tissue of plants, together with pollen competition, would also lead to higher levels of purifying selection acting to remove deleterious variation. Future work should focus on the differential evolution of haploid‐ and diploid‐specific genes to understand the selective dynamics acting on these loci.     

Population sex ratios, sex-specific clonal traits and tradeoffs among these traits in the liverwort Marchantia inflexa

Unisexual, female- or male-biased populations are common among some clonal plants. Within and among populations, the relative frequencies of males and females can be influenced by sex-specific demographic patterns that, in turn, can be a consequence of life-history characteristics. The objectives of this study were to describe (1)  population sex-ratio patterns among habitat patches along a river, and (2)  sex-specific patterns of and correlations among life-history clonal traits including: growth rates, number of meristematic tips and asexual reproduction in the dioecious liverwort, Marchantia inflexa . In Trinidad, a section of a stream was surveyed for the occurrence of female and male sex expressing thalli among habitat patches, and habitat characteristics (canopy openness and patch size) were recorded. The numbers of female and male inflorescences were obtained also. Of the 209 patches observed in the field, 83% did not contain sex expressing thalli, 9% contained thalli of both sexes, and the rest contained only female- (4%) or male-expressing (4%) thalli. Sex expression was less common among small patches, and there was a tendency for sex expression to be less likely among patches with the lowest canopy openness. The proportion of male inflorescences among the bisexual patches ranged from 0.22 to 0.80. In a greenhouse, we planted gemmae (asexual propagules) from field-collected isolates: ten female and eight male. On six occasions, we harvested replicates of each isolate to estimate isolate trait means. Females grew faster, produced more meristematic tips and had lower levels of asexual reproduction than males. Number of meristematic tips was negatively correlated with asexual reproduction and positively correlated with growth rate. These sex-specific patterns may lead to the high frequency of single sex patches as well as biased adult population sex ratios that are observed in the field.     

Life history of Chnoospora implexa (Chnoosporaceae, Phaeophyceae) in culture

The life history of the brown alga Chnoospora implexa J. Agardh (Chnoosporaceae, Scytosiphonales) from Japan was studied in laboratory cultures. This species showed a heteromorphic and diphasic life history, alternating between erect gametophytes and discoid sporophytes. The gametophytes were dioecious and produced isogametes. The zygotes developed into sporophytes at 20°C under long‐day conditions, which formed plurilocular zoidangia. Zoids released from the plurilocular zoidangia developed again into sporophytes that always formed plurilocular zoidangia at 20°C and 25°C in long‐day conditions, and mainly unilocular zoidangia at 25°C in short‐day conditions. Zoids released from unilocular zoidangia developed into dioecious gametophytes. At 15°C zygotic erect thalli were formed and were revealed to be diploid by microspectrofluorometric measurements of nuclear DNA contents. The development and reproduction of unfused gametes were similar to those of zygotes. Some strains showed a direct‐type life history; gametophytic thalli were produced, but not via a sporophytic phase.     

Sex Ratios of Riccia frostii Aust. in Khartoum Province,Sudan

Abstract

Spore dispersal of the dioecious liverwort, Riccia frostii Aust., in Khartoum Province Sudan, is by the Blue Nile in flood. Although a 1 : 1 sex ratio is to be expected, sample counts have shown that females always outnumber males and the actual ratio varies from site to site. Shading has been shown to suppress the development of male thalli, while a pattern analysis and covariance analysis of an apparently homogeneous site showed that male and female thalli reacted differently to microtopographical/substrate differences on the Nile bank. Extreme age of the population produces a change in sex ratio through the earlier death of male thalli; a quickly drying substrate also produces a differential mortality of male thalli. The effect of the high summer temperatures on the dormant spores which remain in situ on the Nile banks is not known but may produce a change in the sex ratio through the differential mortality of the spores.     

INTER-THALLIC SEXUAL REACTIONS IN ACHLYA,A GENUS OF THE AQUATIC FUNGI

Barksdale , Alma Whiffen . (New York Botanical Garden, New York.) Inter-thallic sexual reactions in Achlya, a genus of the aquatic fungi . Amer. Jour. Bot. 47(1): 14—23. Illus. 1960.– Inter-thallic sexual reactions, involving the coöperative induction of sexual organs, have been found to occur between members of homothallic, as well as heterothallic species of the sub-genus Achlya. The induction of sexual organs, both on single thalli and on paired thalli, is temperature-dependent. Sexual organs were initiated at 15°—20°C. in 14 of 27 heterothallic strains that were usually sterile at 25°. The hermaphroditic sexual organs were like those of either A. ambisexualis or A. bisexualis in 19 strains; like those of A. prolifera in 3 strains; like those of A. inflata in 1 strain; and like those of A. americana in 1 strain. In some instances, the sexual reaction between paired thalli was complete at 20°C. but incomplete (i.e., oospores were not formed) at 25°C. The heterothallic and homothallic strains studied could be assigned to 1 of 12 (6 female and 6 male) inter-action types, on the basis of their sexual responses. These sexual types are characterized and their possible usefulness in indicating kinship among species of Achlya is assessed. The possible origin of heterothallic from homothallic forms of Achlya is discussed.     

Interactions in the patterns of vegetative growth and reproduction in woody dioecious plants

Summary Interactions between vegetative growth and reproduction were evaluated in Peumus boldus, Lithraea caustica and Laretia acaulis, three woody dioecious species in central Chile. Phenological observations were made periodically on marked branches of male and female plants, and biomass allocation (dry weight) to vegetative and reproductive tissues was measured. The magnitude of flowering was evaluated in groups of plants in three successive seasons. The patterns of activities are species- and sex-dependent, and cycles of 2–4 years have been established. Branches that produce flowers either do not grow or grow less than branches without flowers, and males and females have differential resource allocation: male branches attain higher biomass values. Groups of plants show seasonal behavior that suggest synchrony in their reproductive activities.     

Cultivation of a hybrid of free-living gametophytes between Undariopsis peterseniana and Undaria pinnatifida: morphological aspects and cultivation period

The kelp Undariopsis peterseniana is warm-water-tolerant, and consequently, there is currently considerable interest in developing commercial cultivation techniques for this species in Korea. U. peterseniana plants have been successfully transferred to the northern coast of Korea beyond their original habitat in Jeju Island (33°30′08.65″N, 126°55′39.02″E). In this study, we cultured a hybrid kelp consisting of a cross between free-living gametophytes of U. peterseniana and U. pinnatifida in an attempt to extend the culture period of Undaria which is an important species for both the abalone industry and for commercial seaweed mariculture for human food applications. Morphological characters and cultivation period were compared between the parent thalli and the hybrid. The cultivation experiment was conducted in Wando, on the southern coast of Korea (34°26′18.68″N, 127°05′43.88″E). The morphological characteristics of the hybrid thalli were intermediate between the two species having shallow pinnated blades and a reduced reproductive organ. Hybrid thalli showed faster growth rates, 1.5 times greater biomass, and a longer cultivation period than the parent thalli. The hybrid strain possessed characteristics that indicate it could be used as an alternative kelp source to supply the abalone feed industry.     

Sex-specific physiology and source-sink relations in the dioecious plant Silene latifolia

Differences in reproductive demands between the sexes of dioecious plants could cause divergence in physiology between the sexes. We found that the reproductive effort of female Silene latifolia plants increased to more than twice that of male plants or female plants that were prevented from setting fruit by lack of pollination after 4 weeks of flowering. Whole-plant source/sink ratios of pollinated females were significantly lower than those of males or unpollinated females because of investment in fruit. We hypothesized that these differences in source/sink ratio between the sexes and within females, depending on pollination, would lead to differences in leaf photosynthetic rates. Within females, we found that photosynthetic capacity was consistent with measurement of whole-plant source/sink ratio. Females that were setting fruit had 30% higher light-saturated photosynthetic rates by 28 days after flowering than females that were not setting fruit. Males, however, had consistently higher photosynthetic rates than females from 10 days after flowering onwards. Males also had approximately twice the dark respiration rates of fruiting females. We found that female reproductive structures are longer-lived and contribute more carbon to their own support than male reproductive structures. Despite the higher rates of leaf dark respiration and lower calyx photosynthetic rates, males fix more carbon than do females. We conclude that females have a sink-regulated mechanism of photosynthesis that allows them to respond to variations in fruit set. This mechanism is not, however, sufficient to explain why male S. latifolia plants have higher rates of photosynthesis, higher source/sink ratios, and lower reproductive allocation, but fail to grow larger than female plants.     

Biosystematics,genetics and upper temperature tolerance ofGigartina teedii (Rhodophyta) from the Atlantic and Mediterranean

Plants ofGigartina teedii from the mediterranean isolated into laboratory culture showedPolysiphonia-type life histories with consistent formation of dioecious gametangial plants, as previously reported for Atlantic isolates. Male and female plants from the Atlantic and Mediterranean were almost completely compatible in terms of cystocarp formation on female plants, and carpospores from positive crosses always formed plants that released viable tetraspores. Sex-linked inheritance of branching pattern was found in all strains, but showed varying degrees of expression. Female plants were more branched than male plants and it is suggested that this may be an adaptation for spermatial capture.G. teedii plants showed differences in morphology in culture that are considered to be genetically-based. Preliminary studies of tip elongation showed that Mediterranean strains may have up to three times the elongation rates of Atlantic strains at 15°C, . Such genetic variation in fully-interbreeding strains suggests that populations of this species in the Atlantic and Mediterranean are genecodemic. All strains showed an upper temperature tolerance of 31°C when tested at 1°C intervals from 29—34°C. An upper temperature tolerance of 31–32°C was found for the related speciesG. intermedia from Korea and Japan, butG. johnstonii from the Gulf of California showed an upper tolerance of 32–33°C.     

Eine diözische Flechte

The lichenLecidea verruca, growing parasitically on other crustaceous lichens, is, contrary to most other lichenized ascomycetes, dioecious. The male thalli bearing spermogonia are mostly smaller than the female ones.

     

Sex‐specific interaction between arbuscular mycorrhizal and dark septate fungi in the dioecious plant Antennaria dioica (Asteraceae)

Male and female plants of dioecious species often differ in their resource demands and this has been linked to secondary sexual dimorphism, including sex‐specific interactions with other organisms such as herbivores and pollinators. However, little is known about the interaction between dioecious plants and fungal root endophytes. Plants may be simultaneously colonised by arbuscular mycorrhizal (AM) and dark septate (DS) fungi. While it is well established that AM mutualism involves reciprocal transfer of photosynthates and mineral nutrients between roots of host plants and these fungi, the role of DS fungi remains controversial. Here, we report the temporal and spatial variation in AM and DS fungi in female, male and non‐reproductive Antennaria dioica plants in three natural populations in Finland during flowering and after seed production. Females had higher colonisation by AM fungi, but lower colonisation by DS fungi than male and non‐reproductive plants. The higher AM colonisation was observed during flowering, and this difference varied among populations. Our results suggest that females and males of A. dioica interact with AM and DS fungi differently and that this relationship is dependent on soil fertility.     

Effects of photoperiod on the reproductive condition of Nile grass rats (Arvicanthis niloticus) from an equatorial population

We evaluated the effects of photoperiod on the reproductive condition of male and female Nile grass rats (Arvicanthis niloticus) descended from members of an equatorial population trapped 2°S of the equator. Study animals housed in 12:12 light:dark (LD) cycles were transferred either to short photoperiod (9:15) or long photoperiod (15:9) for 9 weeks (males) or 11 weeks (females), and various reproductive parameters were assessed. We observed no differences between short‐ and long‐day males with respect to plasma concentration of testosterone, testicular mass, seminal vesicle mass, or spermatogenesis. Similarly, we observed no differences between short‐ and long‐day females with respect to oestrous cycles, uterine mass, follicle size, or presence of corpora lutea. Reproductive parameters of male and female A. niloticus housed in short‐ and long photoperiods were similar to those typically observed among animals descended from the same equatorial population and housed in LD 12:12. Thus, photoperiod appears not to elicit changes in reproductive condition among A. niloticus from populations whose native habitat lies within 2° of the equator. These data contrast with the results of other studies indicating that photoperiod alters reproductive condition in A. niloticus populations living >10° from the equator.     

Impacts of floral gender and whole-plant gender on floral evolution in Ecballium elaterium (Cucurbitaceae)

Investigation of gender specialization in plants has led to several theories on the evolution of sexual dimorphism: reproductive compensation, based on enhanced reproductive efficiency with gender specialization (flowers should be larger on dioecious plants); Bateman's Principle, based on sex-specific selection (display for pollinator attraction in males and seed set in females); and intersexual floral mimicry, based on mimicry of a reward-providing gender by a non-reward providing gender (reduced dimorphism in dioecious plants due to increased spatial separation of male and female flowers). These theories were evaluated in Ecballium elaterium, which contains two subspecies, elaterium (monoecious) and dioicum (dioecious). Our results show that flowers of the dioecious subspecies are larger and allocate more to reproductive organs than do flowers of the monoecious subspecies. Both subspecies are sexually dimorphic (male flowers larger than female flowers). Variance in flower size among populations is greater in the dioecious subspecies. Finally, there is sufficient genetic variation to enable ongoing response to selection; genetic correlation constraints on independent response of female and male flowers may be stronger in the monoecious subspecies. Our findings provide support for aspects of all three theories, suggesting that the evolution of floral dimorphism is based on a complex interplay of factors.     

LIFE HISTORY AND HYBRIDIZATION STUDIES ON GIGARTINA STELLATA AND PETROCELIS CRUENTA (RHODOPHYTA) IN THE NORTH ATLANTIC1

Fourteen isolates of the crustose marine red alga Petrocelis cruenta J. Agardh from various localities in the British Isles, France (including the type locality), Spain and Portugal gave rise in culture to dioecious foliose plants identifiable as Gigartina stellata (Stackhouse) Batters although two isolates formed only sterile foliose blades. A total of 145 isolates of Gigartina stellata were also grown in culture from various localities in the U.S.A. (Maine), the British Isles, Iceland, Denmark, France, Spain and Portugal using both carpospores and vegetative blade apices. Two basic types of life history were found among these isolates: a direct-type life history involving the formation of further foliose plants from carpospores, some isolates of which also form spermatangia on the same papillae as the cystocarps; and a heteromorphic-type in which only crustose plants resembling Petrocelis cruenta are formed from carpospores. Only heteromorphic-type life histories were found from Spain and Portugal. Both life history types were found in plants from the U.S.A., the British Isles and northern France. Only direct-type life histories were found in plants from Iceland and Denmark. Some Petrocelis-like crusts derived from field collected G. stellata carpospores and Petrocelis crusts of hybrid progeny formed tetrasporangia in 8:16 h LD, 10° C but not in 8:16 h LD, 15° C; 16:8 h LD 10° C or 15° C; and 10:6.5:1: 6.5 h LDLD, 10° C. The spores thus formed were viable and produced normal dioecious male and female gametophytes. Short day and low temperature conditions appear necessary for tetrasporogenesis. The results from crossing experiments with 32 male and 27 female isolates of the heteromorphic-type derived from both G. stellata and P. cruenta showed that two virtually non-interbreeding populations with a high degree of geographical separation exist in the north-eastern Atlantic. Morphological differences between plants from each population are described. On the basis of culture and crossing results, Petrocelis cruenta J. Agardh is placed in synonymy with Gigartina stellata (Stackhouse in Withering) Batters.