Examination of the ability of gonadal sex change in primary males of the diandric wrasses Halichoeres poecilopterus and Halichoeres tenuispinis: estrogen implantation experiments

Primary males that function as males throughout their lives are often found among protogynous fishes such as wrasses and parrotfishes. However, the issue of whether the sexuality of primary males involves gonochorism or hermaphroditism remains uncertain. To clarify this, we implanted estradiol-17beta (E2) into the body cavities of primary males of two protogynous wrasses, Halichoeres poecilopterus and Halichoeres tenuispinis. At 51-63 days after implantation, primary males with E2 treatment in both wrasses were observed to develop ovarian tissues. These results suggest that primary males of Halichoeres wrasses potentially have the ability to change sex and that estradiol-17beta is related to gonadal transitions in primary males.     

New aspects of sex change among reef fishes: recent studies in Japan

New aspects of sex change in reef fishes are reviewed with special emphasis on recent studies in Japan. For protogyny, studies on both monandric and diandric species have been conducted, but the distinction of primary males from prematurational secondary males seems to need further examination. For protandry, detailed field studies on anemonefishes have revealed alternative life-history styles associated with movements between hosts before or after maturation. The most interesting new aspect has been the discovery of 2-way sex change within a species. Conditions for evolution of 2-way sex change are examined in relation to the size-advantage model and social control mechanisms. A fish may change sex when it becomes dominant in a mating group, but a dominant fish may also change sex in the reverse direction when its social status changes to subordinate through inter-group movement. Two-way sex change has hitherto been reported only from basically protogynous fishes (e.g., Gobiidae, Pomacanthidae, Cirrhitidae, Epinephelinae). Possibilities of the reverse sex change in the protandrous anemonefishes are discussed with data from some unpublished studies.     

Reproductive cycle and sex inversion in razor fish, a protogynous labrid in the southern Mediterranean Sea

The reproductive biology of the Mediterranean razor fish Xyrichthys novacula was investigated by demographic data and histological analysis of the female, intersexual and male gonads. Specimens were collected by bottom trawl on a monthly basis between June 2000 and July 2001 in a sandy bay in southern Thyrrenian. Gonad histology confirmed that the Mediterranean razor fish is a monandric, protogynous hermaphrodite. Females reached first sexual maturity at 100 mm ( L _T) and the estimated mean L _T at first maturity ( L ₅₀) was 125 mm. Females exhibited asynchronous ovarian development and multiple ovulations occurred over the spawning period. Vitellogenesis started in early May and spawning occurred from late May until late September. Sexual transition involved a large‐scale atresia of all oocyte stages and a massive degeneration of ovarian tissue followed by primordial germ cells proliferation. Sex change began at spawning time (June) but transitional individuals tended to cluster at the end of the reproductive period (September). They accounted for 17·1% of the population sampled and were found in a broad size range (105–150 mm L _T).     

Testing a new version of the size-advantage hypothesis for sex change: sperm competition and size-skew effects in the bucktooth parrotfish, Sparisoma radians

A variety of field studies suggest that sex change in animalsmay be more complicated than originally depicted by the size-advantagehypothesis. A modification of the size-advantage hypothesis,the expected reproductive success threshold model, proposesthat sperm competition and size-fecundity skew can stronglyaffect reproductive pay-offs. Size-fecundity skew occurs ifa large female's fecundity is markedly higher than the aggregateof the other members of her social group and, together withpaternity dilution from sperm competition, can produce situationsin which large females benefit by deferring sex change to smallerfemales. Deferral by large females can create sex-size distributionscharacterized by the presence of large females and small sex-changedmales, and it is precisely these distributions that the traditionalsize-advantage model cannot explain. We tested the predictionsof the new model with the bucktooth parrotfish, Sparisoma radians,on coral reefs in St. Croix, U.S. Virgin Islands. Collectionsand spawning observations determined that the local environmentalregime of S. radians is characterized by pervasive sperm competition(accompanying 30% of spawns) and factors that can produce substantialsize-fecundity skew in social groups. Dominant male removalexperiments demonstrate that the largest females in social groupsoften do not change sex when provided an opportunity. Instead,smaller, lower-ranking females change sex when a harem vacancyarises. This pattern of sex change is in contrast to virtuallyall previous studies of social control of sex change in fishes,but provides strong support for the general predictions of theexpected reproductive success threshold model.     

Group spawning results from the streaking of small males into a sneaking pair: male alternative reproductive tactics in the threespot wrasse Halichoeres trimaculatus

In many protogynous wrasses, large males with bright coloration (terminal phase males, TP males) establish mating territories and pair-spawn with females. In contrast, small primary males with drab coloration (initial phase males, IP males) are non-territorial and adopt three alternative reproductive tactics—group spawning, streaking, and sneaking. We investigated how IP males of the threespot wrasse Halichoeres trimaculatus use these tactics in different situations. The mating frequency of the IP males was positively correlated with their courtship frequency, but not with their body size. Larger IP males tended to attack the smaller ones at the mating sites. This indicates that the larger IP males attempted to exclude the smaller ones from mating with the intention of minimizing the number of IP males involved in group spawning and ultimately leading to pair spawning (sneaking). However, the larger IP males were unable to completely exclude the smaller males because the intensity of the attack by the larger IP males was weak. Consequently, the smaller IP males could easily streak into the sneaking of larger IP males, thereby resulting in group spawning.     

Density-dependent protogynous sex change in territorial-haremic fishes: models and evidence

Several hypotheses of the proximate control of protogynous (female-to-male)sex change propose that social group composition triggers sexchange, but they do not address how proximate cues are alteredby population density. I present three mutually exclusive encounter-ratethreshold hypotheses that assume that population density determinesrates of contact between social group members and that ratesof contact are cues for sex change. Different densities arepredicted to induce sex change, depending on the encountersassumed to be important in the sex change process (e.g., encounterswith smaller and larger individuals). Tests of the models usea pomacanthid angelfish(Centropyge potten) to show that continuedpresence of a smaller (female) conspecific is needed for sexchange, and that continued presence of a larger (male) conspecificcan either inhibit sex change or prevent its behavioral stimulation.Using constant social group composition, sex change is preventedat higher density but not at a lower density. The absolute encounter-ratethreshold hypothesis, which predicts sex change under intermediate-densityconditions, is the most probable model of the social controlof sex change in C.potteri     

Sex structure and potential female fecundity in a Epinephelus guttatus spawning aggregation: applying ultrasonic imaging

Ultrasonic imaging was used to determine the spawning population structure and develop a fecundity estimation model for a red hind Epinephelus guttatus spawning aggregation within the Red Hind Bank Marine Conservation District, St Thomas, U.S.V.I. The spawning population showed considerable within‐month and between‐month variation in population size‐ and sex‐structure. In the spawning season studied, males appeared to arrive at the aggregation site first in December although females represented a large proportion of the catch early in the aggregation periods in January and February. Spawning occurred in January and February, and size frequency distributions suggested that an influx of small females occurred during the second spawning month. An overall sex ratio of 2·9 : 1 (female : male) was recorded for the whole reproductive season. The sex ratio, however, differed between months and days within months. More females per male were recorded in January than in February when the sex ratio was male biased. Fecundity estimates for this species predicted very high potential fecundities (2·4 × 10⁵−2·4 × 10⁶ oocytes). The ultrasound model also illustrated a rapid increase in potential female fecundity with total length. Ultrasonic imaging may prove a valuable tool in population assessment for many species and locations in which invasive fishing methods are clearly undesirable.     

Social control of terminal phase transition in primary males of the diandric wrasse, <Emphasis Type="Italic">Halichoeres poecilopterus</Emphasis> (Pisces: Labridae)

In many diandric fishes, large territorial males with bright body coloration (terminal phase (TP) males) are derived either from initial phase (IP) females that change sex to male or from IP primary males that change color and behavior, but do not change sex. The mechanism controlling the transition of IP primary males into TP males is not well understood. We conducted cohabitation experiments to examine social conditions favoring TP transition by primary males in the diandric wrasse, Halichoeres poecilopterus. IP primary males always started TP-specific sexual behavior in the presence of a smaller subordinate, and subsequently acquired TP body coloration. In contrast, primary males under subordinate conditions often performed female-like sexual behavior. In pairs with similar body sizes, both individuals initiated TP male behavior. The results suggest that TP transition in primary males may be closely related to a dominance relationship (or size order) within social groups, as it is in the case of sex change by females.     

Rapid larval growth predisposes sex change and sexual size dimorphism in a protogynous hermaphrodite, Parapercis snyderi Jordan & Starks 1905

The temporal relationship between growth history, sex-specific growth divergence and sex change was investigated in the haremic sandperch Parapercis snyderi using otolith microstructure and gonad histology. Parapercis synderi was found to display rapid near-linear growth with a maximum longevity of 303 days. All individuals matured first as female and later changed sex to become male (monandric protogynous hermaphroditism). Individual age-based growth histories obtained from otolith increment widths illustrated that males were larger than females at any given age. Males were found to diverge from the female growth trajectory during two ontogenetic periods; during the larval period and during the period that sex change took place. In addition, male otoliths contained a discontinuity, or 'check mark', associated with the rapid increase in otolith growth during the sex-change period. This microstructural feature was absent from all female otoliths. Accelerated growth in male otoliths lasted up to 25 days, following check-mark formation, after which time otolith growth returned to the pre-check-mark rate. Given the isometric relationship between otolith and somatic growth in P. synderi , and the temporal relationship between the time of check-mark formation and gonad condition, these results strongly suggest that individuals accelerate somatic growth during sex change to become the largest members of the population. Moreover, evidence suggests that the factors that determine the initial growth of larvae influence which individuals will later become males and achieve the highest reproductive success.     

Sex ratio, life-history invariants, and patterns of sex change in a family of protandrous gastropods

Application of optimality theory to the evolution of life histories has been broadly successful in predicting the conditions favoring sex change, the type of change, and the timing of such changes. The size advantage hypothesis predicts that the optimal size at which an individual should change sex is a function of its size and the size and sex of its potential mates. I collected data on the size, sex, and grouping of individuals of 27 populations of 19 species of the calyptraeids, a family of protandrous marine gastropods that includes Crepidula. These data are used to test the following predictions about variation in size at sex change: (1) sex ratio is biased toward the first sex; (2) the ratio of the size at sex change to the maximum size is a life-history invariant; and (3) species that form variable-sized groups or stacks have more variation in size at sex change than species that show less variation in stack formation. Across all 19 species, sex ratio was not significantly more often biased toward the first sex than it was toward the second sex, although sex ratios were significantly male biased more often than they were significantly female biased. Sex ratios ranged from 0.05 to 0.91, and this variation was related to mode of development, skew in size distribution, and frequency of stacking, but not with maximum body size. There was little evidence that the ratio of size at sex change and maximum size is invariant. There is evidence that one of the main underlying assumptions of this life-history invariant, that male fertility increases with the same function of size in all species, is invalid for calyptraeids and probably for other animals. Finally, species that form larger stacks or mating groups had more variation in size at sex change within a population than species that were generally solitary. These results suggest that information about individual groupings should be included in predictions of life-history theory and that more information about the relationship between male fitness and size is also needed.     

Sex change in plants and animals: a unified perspective

The capacity of organisms to change their sex has evolved independently in several plant and animal lineages. Sex change has been widely studied, but research approaches have differed for plants and animals, and conclusions have often been taxon‐specific. Although sex allocation theory provides a unifying framework for the study of sex change, this unity has not always been appreciated, especially in the botanical literature. Here, we review sex change with regard to its representation in relation to taxonomy and other sexual systems, with regard to its suggested adaptive benefits, and to the role of taxon‐specific body architecture, such as modularity and gonadal structure. We highlight differences and similarities between plants and animals and suggest promising lines of future research.     

Presence of primary and secondary males in a population of the protogynous Synbranchus marmoratus

Synbranchus marmoratus , the 'swamp eel', is a protogynous diandric fish. The primary and secondary males can be distinguished from each other easily by differences in gonadal morphology. Primary males have lobular, unrestricted testes with central efferent ducts. The secondary male has a 'lamellar' testis, efferent ducts are present in the ventral region (new formation), lateral supports and it is covered by the former ovarian capsule. The length of primary male varies from 13 to 88 cm while secondary males range from 56 to 91 cm. Transitional individuals vary between 45 and 60 cm in length. The swamp eel population studied is composed of 80% primary males and 20% secondary males. Although gonadosomatic indices are always higher in primary males, they increase in secondary ones as the newly acquired stage progresses. The lamellar organization of the gonad of secondary males, the absence of this kind of testis in individuals smaller than 56 cm, and the presence of hermaphroditic fish, is evidence for protogyny in this species.     

2000-2020年祁连山植被净初级生产力时空变化及其驱动因素

植被净初级生产力(NPP)是评价植被生长的重要参数,也是评估陆地生态系统质量与功能的重要指标。基于MODIS NPP、数字高程模型(DEM)、气象水文及人类活动数据,采用空间分析、趋势分析,分别从像元尺度和县域尺度识别了2000-2020年以来祁连山NPP时空变化特征,采用偏相关分析研究了NPP对年均温和年降水的响应,并借助地理探测器模型揭示了NPP变化的驱动因素,最后采用Hurst指数预测了NPP未来变化趋势。结果表明:2000-2020年祁连山平均NPP呈波动增加趋势,年均增加2.38 g C/m²,其中栽培植被和阔叶林增长最为明显。近20年,像元尺度上有75.37%的区域NPP增加,主要位于东南部;县域尺度上,古浪、平安、化隆和永登县NPP增速较快,而祁连、海西、德令哈和门源县增速较慢。祁连山NPP空间分布具有明显的集聚性,高值集聚区主要位于东南部,而低值集聚区主要位于西北部。年均温和降水量的增加均促进了NPP的增加,但不同区域NPP对气温和降水的响应有明显差异。降水量、饱和水气压差和蒸散发是NPP变化的主要驱动因子,驱动因子之间对植被NPP变化存在交互作用,分为双因子增强和非线性增强效应。未来祁连山NPP变化以增加非持续性为主,说明植被变化面临较大不确定性。研究结果有助于揭示全球气候变化背景下区域植被NPP对气候变化及人类活动的非线性响应机制,亦可为祁连山生态保护与可持续发展提供理论依据。     

2000-2008年中国东北地区植被净初级生产力的模拟及季节变化

利用GLOPEM-CEVSA模型模拟并分析了中国东北地区2000-2008年植被净初级生产力(NPP)时空分布格局及其影响因素,并以4个森林生态站点(大兴安岭、老爷岭、凉水和长白山森林生态站)为例研究了东北地区森林NPP季节变化特征及其环境驱动.结果表明:2000-2008年,东北地区植被年均NPP为445 g C·m-2·a-1;整个研究区沿长白山山脉到小兴安岭山脉地区以及三江平原部分地区的NPP最高,沿长白山山脉到小兴安岭山脉西侧的辽河平原、松嫩平原东部、三江平原和大兴安岭地区次之,西部稀疏草原和荒漠地区的NPP最低.东北地区森林生态系统年均NPP最高,其次为灌丛、农田和草地,荒漠最低.森林生态系统中,针阔混交林年均NPP最大(722 g C·m-2·a-1),落叶针叶林年均NPP最小(451 g C·m-2·a-1).研究期间,森林NPP无显著年际变化,其中2007、2008年较往年NPP大幅增加,很可能与该地区期间气温上升有关(较往年偏高1 ℃=～2℃).东北地区森林自北向南生长季开始时间逐渐提前,生长季变长.     

Sex ratio and male sexual characters in a population of blue tits, Parus caeruleus

Sex allocation theory proposes that parents should bias thesex ratio of their offspring if the reproductive value of onesex is greater than that of the other. In the monogamous bluetit (Parus caeruleus), males have a greater variance in reproductivesuccess than females, and high-quality males have higher reproductivesuccess than high-quality females due to extrapair paternity.Consequently, females mating with attractive males are expectedto produce broods biased toward sons, as sons benefit more thandaughters from inheriting their father's characteristics. Songand plumage color in birds are secondary sexual characters indicatingmale quality and involved in female choice. We used these malesexual traits in blue tits to investigate adaptive sex ratiomanipulation by females. We did not find any relationship betweenmale color ornamentation and brood sex ratio, contrary to previousstudies. On the other hand, the length of the strophe bout (i.e.,the mean number of strophes per strophe bout) of fathers waspositively related with the proportion of sons in their broods.The length of the strophe bout is supposed to reflect male qualityin terms of neuromuscular performance. We further showed thatsons produced in experimentally enlarged broods had shorterstrophe bouts than sons raised in reduced broods. These resultsare consistent with the hypothesis that females adjust the sexratio of their broods in response to the phenotype of theirmate.     

Sex allocation and larval competition in a superparasitizing solitary egg parasitoid: competing strategies for an optimal sex ratio

1. Parasitic Hymenoptera reproduce by arrhenotokous parthenogenesis, and females of these species are able to control their progeny sex ratios. In structured populations of parasitic Hymenoptera, primary sex ratios are often highly biased toward females. However, sex ratio can be adjusted to the quality of encountered patches or hosts or be modified by differential developmental mortality.
2. In this paper, the effects were evaluated of the quality of encountered hosts and developmental mortality on the sex ratio in Anaphes victus , a solitary egg parasitoid whose first instar larvae present a sexual dimorphism and where superparasitism is regulated by larval fights between first instar larvae.
3. The results showed that a female-biased sex ratio is allocated to unparasitized hosts. In the presence of parasitized hosts, the second (superparasitizing) female produced a significantly higher sex ratio than the first female but the tertiary sex ratio (sex ratio at emergence) was not significantly different from the sex ratio produced with unparasitized hosts. The increase in the primary sex ratio produced by the second female was mostly compensated by the higher mortality of male larvae.     

Responses of vegetation structure and primary production of a forest transect in eastern China to global change

Aim A regional model of vegetation dynamics was enhanced to include biogeochemical cycling of nitrogen and was then applied to a forest transect in east China (FTEC) in order to investigate the responses of the transect to possible global change. Location Eastern China. Methods Biomass and nitrogen concentration of green and nongreen portions of vegetation, moisture contents of three soil layers, and total and available soil nitrogen are included as state variables in the enhanced model. The model was parameterized and validated against field observations of biomass, productivity, plant and soil nitrogen concentration, nitrogen uptake, a vegetation index derived from satellite remote sensing and digital maps of vegetation and soil distributions along a forest transect in eastern China (FTEC). The model was applied to FTEC in order to investigate the responsive characteristics of the ecosystems to global climatic change. Scenarios of climate change under doubled CO₂ produced by seven general circulation models (GCM) were used to drive the model. Results The simulations indicated that the model is capable of simulating accurately potential vegetation distribution and net primary productivity under contemporary climatic conditions. The simulations for GCM‐projected future climate scenarios with doubled atmospheric CO₂ concentration predicted that broadleaf forests would increase, but conifer forests, shrubs and grasses would decrease; and that deciduous forests would have the largest relative increase, but evergreen shrubs would have the largest decrease. Conclusions The overall effects of doubling CO₂ and climatic changes on FTEC were to produce an increased net primary productivity (NPP) at equilibrium for all seven GCM scenarios. The inclusion of nitrogen dynamics in the model imposes more constraint on the responses of FTEC to climatic change than the previous version of the model without nitrogen dynamics. Temperature exerts a stronger control on NPP than precipitation, as indicated by the negative correlations between NPP and temperature. The southern portion of FTEC, at latitudes less than 33 °N, show much larger increases in annual NPP than in the north. However, the predicted range of NPP increases is much larger in the north than in the south.     

气候变化和人类活动对中国陆地生态系统总初级生产力的影响厘定研究

总初级生产力(GPP)是生态系统植被光合作用生成有机物的能力表征,是生态系统服务功能的基础,关系到区域社会经济可持续发展及区域生态安全。基于生态系统过程模型CEVSA2,应用中分辨率成像光谱仪(MODIS)卫星遥感的叶面积指数数据产品(MCD15A2H),以强迫法构建了遥感数据驱动的模型新版本——CEVSA-RS;基于CEVSA-RS模拟分析了气候变化和人类活动对中国陆地生态系统GPP时空变化的相对影响,从气候潜在总初级生产力(GPP_CL)和现实总初级生产力(GPP_RS)的大小和趋势两方面厘定了人类活动影响。2000至2017年全国平均潜在GPP(1016.36 gC m^-2a^-1)略高于对应现实GPP(962.85 gC m^-2a^-1),但存在明显的空间分异:长江以南大部、秦岭、太行山脉以东以及大兴安岭以东和长白山地区等森林植被覆盖区,现实GPP高于潜在GPP;而西部草地及灌丛等地区现实GPP低于潜在GPP。全国GPP呈显著增加趋势(P<0.05)...