Sex Expression and the Evolutionary Advantages of Male Flowers in an Andromonoecious Species, Sagittaria guyanensis subsp. lappula (Alismataceae)

Sex expression in Sagittaria guyanensis H.B.K. subsp. lappula (D. Don) Bojin is typically andromonoecious while the other species in the same genus are basically monoecious. The evolutionary advantages of male flowers and hermaphrodite flowers in S. guyanensis subsp. lappula were assessed by measuring sex allocation and pollen movements in two wild populations of the species. Two cultivated populations served as controls. The percentage of male flowers was very low in the two wild populations in Dongxiang, Jiangxi Province and Wuyishan, Fujian Province, viz., 2.48% and 0.96% respectively. In the two cultivated populations, male flower percentage significantly increased when the soil was of higher nutrient content. This indicates that the allocation to male versus female reproduction might change in response to environmental factors. Pollen production per male flower was 4.1 times higher than that of a hermaphrodite flower. The floral shape and size of male and hermaphrodite flowers were similar. No difference was observed between these dimorphological flowers in pollen germination rate in vitro and in the speed of pollen tube growth in vivo . Anthesis was only 4-5 h. Male flowers usually opened 0.5 h earlier than hermaphrodite flowers. An unexpected finding was that no pollen from the male flowers was found on the stigmas of the hermaphrodite flowers, in spite of the occasional visits by insects to both types of flowers in both wild and control populations. A consistent pattern of fruit development was found to exist in open pollinated flowers as well as in flowers that had been bagged. The sex ratios did not have significant influence on fruit set. Approximately 25% of the pistils in a gynoecium failed to develop into fruits because no pollen was deposited on them, indicating that the fruit set of this andromonoecious plant is mainly affected by pollen limitation rather than resource limitation. Reproduction in S. guyanensis subsp. lappula in the habitats was dependent on self_pollination in hermaphrodite flowers. The male flowers in this species might be a potential source of additional pollination and may facilitate cross_pollination. The fact that the flowers of monoecious species in Sagittaria pollinated by a wide diversity of insect visitors may contribute enormously to the diversification of sex expression in this genus.

High-density genetic mapping identified a major locus for environmental sex expression in pumpkin(Cucurbita moschata Duch.)

Long-day length and high temperature inhibit sex expression in pumpkin(Cucurbita moschata Duch.),and therefore directly impact the production potential.In this study,female flowering patterns in photoperiod-insensitive(PPIS)and photoperiod-sensitive(PPS)germplasms differed significantly in a moderately long day and high temperature environment.However,both germplasms exhibited a similar response in short day with either low temperature or high temperature environment.Photoperiod sensitivity led to this difference in sex expression between the germplasms.For the traits of 1st female flowering node(FFFN)and number of female flowers(NFF),high-density linkage map construction and quantitative trait locus(QTL)mapping were performed using SLAF-seq technology and 162 F_(2) individuals generated from PPIS and PPS.In total,4655 SLAFs were selected and mapped on 20 linkage groups(LGs).The total map length was 2502.01 cM with an average interval distance of 0.75 cM.Major QTLs for both FFFN and NFF were detected on LG6 with intervals of 7.89 and 17.67 cM and PVE values of 30.5%and 22.9%,respectively.Further analyses of the major locus for FFFN revealed 73 protein-coding genes.Among them,4 were related to sex expression,photoperiod flowering,and hormone response.An InDel(insertion and deletion)marker partially correlated with FFFN of the F_(2) population was also developed.Our study identified the QTL for the sex expression response to environmental factors using the high-density linkage map.The identified candidate genes and markers will provide useful information about the molecular interaction between the environment and sex expression and for marker-assisted selection of pumpkin environment-insensitive resources.

Modulation of Sex Expression in Four Forms of Watermelon by Gibberellin, Ethephone and Silver Nitrate

There has been no systematic research on the effect of plant growth regulators and silver nitrate treatments on the control of sex expression in watermelon. In this study, we tested sex expression responses of four watermelon forms(monoecism, gynoecism, andromonoecism,and hermaphrodite) to gibberellin, ethephon and silver nitrate treatments. Results have shown that, in monoecious plants, gibberellins(GA_3)and ethephon treatments reduced the percentage of female flowers and delayed the occurrence of the first female flower, while silver nitrate induced the formation of bisexual flowers. In gynoecious plants, both ethephon and silver nitrate treatments transformed some female flowers into bisexual flowers, and treatment with ethephon resulted in a mass of abnormal flowers, while no obvious effect of treatment with GA_3 was observed. In andromonoecious plants, ethephon and GA_3 treatments delayed the occurrence of the first bisexual flower, and GA_3 reduced the percentage of bisexual flowers, while no distinct effect for silver nitrate treatment was observed. In hermaphroditic plants, ethephon treatment induced the appearance of numerous abnormal flowers, while no obvious effects for GA_3 and silver nitrate treatments were observed.We analyzed the transcription levels of all the expressed aminocyclopropane-1-carboxylic acid synthase(ACS) homologues in two gynoecious mutants and their wild types. We also tested the gene expression of CitACS4 which had been recognized as the andromonoecious gene in all treatments. All these results suggested that the best masculinizing treatment for breeding of the gynoecious line is silver nitrate, which repressed the expression of CitACS4 and induced many bisexual flowers for use in self-fertilization subsequently.

《沉沦》艺术美新论

由小说内容表现出的青春期性行为反应,探讨了郁达夫的小说《沉沦》的审美教育作用;从叙述艺术视角分析了作品的诗化小说特征;从作品的思想及创作手法上考察了作品的现代主义思想和表现主义艺术特征;并从背景、环境等方面分析了小说特色的成因。

Gender plasticity uncovers multiple sexual morphs in natural populations of Cedrus deodara (Roxb.) G. Don

Background:The expression of gender in gymnosperms is challenging because the extent of variability in gender segregations in tree crowns and selection pressure of nature can modify the gender through time.Methods:An in-depth investigation on spatial segregation of genders in tree crowns and sex expression of a total of 500 trees in five different natural populations of Cedrus deodara was carried out and verified the occurrence of subdioecious(co-existence of male,female,and monoecious)genders in C.deodara.Results:Five different sexual morphs were apparent among the 500 selected individuals as(1)pure male(M):bearing only male strobili in the whole crown with 22.2%;(2)pure female(F):bearing only female strobili in the whole crown with 12.4%;(3)mixed-monoecious(MM):bearing both male and female strobili with 13.6%;(4)predominantly female(PF)with 25.6%;and(5)predominantly male(PM)with 26.2%.The occurrence of multiple sexual morphs resulted from the complex selection pressure,which increased the stability of the populations.The segregation of genders in crown layers deemed to increase the fitness that may be a mechanism for accelerating outcrossing.The results of the study suggest that the subdioecious gender expression in C.deodara is evolved through the monoecy–paradioecy pathway.The production of male strobili revealed non-significant effect of tree sizes whereas a significant effect was observed for the production of female strobili.Our study established that the total pollen and seed outputs in C.deodara changed frequently according to gender expression.Conclusions:The size of the tree,resource availability,sex allocation,plant architecture,gender segregation in crowns,and habitat conditions are the prominent causes for gender plasticity.