白色霞水母生活史的实验室观察

本文首次描述了白色霞水母从受精卵至碟状体的生活史。(1)包括受精卵、卵裂、囊胚以及浮浪幼虫等在内的胚胎发育各期均在开放的水体中,在20·8 -21·4℃浮浪幼虫于受精后14 h出现; (2)浮浪幼虫在定置前形成一种凸面的圆形浮浪幼体囊,除了浮浪体囊外,螅状体还可产生足囊和通过产生匍匐茎形成囊胞进而发育成新的螅状体; (3)尽管偶而产生2个碟状体但仍为典型的单碟型横裂; (4)新释放的碟状幼体绝大多数为8个缘叶, 8个感觉棍和8对钝圆的缘瓣,但畸形个体最多12个,最少6个缘叶; (5)雌雄个体间的交互作用对产卵和受精是非常重要的因子[动物学报52 (2) : 389 -395 , 2006]。     

盐度对沙蜇有性繁殖阶段早期发育的影响

近几十年来,沙蜇的频繁暴发给东亚海域的海洋生态系统带来了广泛影响。在秋季,沙蜇成熟的雌雄水母体在沿岸水域聚集产卵,有性繁殖产生的受精卵发育成新的底栖螅状体,为螅状体种群数量进行补充。河口浅滩海域为沙蜇的繁育地,沿岸盐度较低,在秋季降雨期盐度多变,较低、多变的盐度可能对沙蜇有性繁殖阶段的早期发育产生重要作用,从而影响螅状体种群数量的补充。实验设置了4种不同盐度(15、20、25、30)试验组,在不同盐度下对沙蜇受精卵进行培养,探讨盐度对沙蜇早期发育过程中受精卵、浮浪幼虫发育以及早期螅状体生长及存活的影响。试验结果:沙蜇受精卵胚胎发育的适宜盐度为20,发育基本与盐度25、30同步,盐度15受精卵细胞发育迟缓,发育率显著降低;浮浪幼虫发育适宜盐度为20和25,两组浮浪幼虫附着变态率高于盐度15、30,盐度15时浮浪幼虫活力明显降低、发育迟缓,浮浪幼虫在盐度15时水中存活时间较长可达8 d,但附着时间集中在培养后的3、4天,与其他组相同;早期螅状体幼体适宜盐度为20、25、30,早期螅状体存活率、相对增长率及特定生长率均显著高于盐度15,三组间差异不显著。结果表明,盐度显著影响沙蜇有性繁殖阶段的早期发育,随着受精卵至螅状体的发育生长,其对盐度的适应范围逐步扩大。     

巴布亚硝水母无性生殖过程及形态变化北大核心CSCD

以巴布亚硝水母(Mastigias papua)为研究对象,观察了水螅体无性生殖产生类浮浪幼虫胞芽、类浮浪幼虫胞芽变态发育为水螅体、水螅体横裂产生碟状体以及碟状体发育为水母体的过程和形态变化。25℃时类浮浪幼虫胞芽经过93 h后变态发育为水螅体。换用带有虫黄藻的天然海水并将培养温度从25℃升至27℃后,水螅体开始横裂,萼部触手环下方产生缢痕。水螅体横裂开始47 h后,缢痕更加明显,其上方部分发育为碟状幼体,后期碟状体搏动越来越频繁,最后释放。释放后的碟状体在实验室培养条件下21 d后可发育为水母幼体。巴布亚硝水母虽然只有产生类浮浪幼虫胞芽一种无性生殖方式,但繁殖速度较快,27℃时1个水螅体平均1 d可产生1.7个类浮浪幼虫胞芽,类浮浪幼虫胞芽可在3或4 d的短时间内附着变态。     

海蜇四触手螅状体两种发生方式的观察

通过对海蜇(Rhopilema esculentum)胚胎发育和足囊萌发进行连续观察,发现海蜇四触手螅状体存在2种形成方式:第一种方式为海蜇受精卵在水温(26±1)℃条件下经6 h发育到浮浪幼虫期,浮浪幼虫25 h后经过杯状体阶段,变态发育成四触手螅状体,进入附着生活阶段;第二种方式为足囊在适宜条件下萌发,首先形成杯状...     

盐度对海蛰各发育阶段幼体的影响——兼论辽东湾海蜇资源锐减的原因

在室内控制条件下,盐度为2—31.42‰范围内,本文就盐度对海蜇(Rhopilema esculenta Kishinouye)浮浪幼虫、螅状幼体、碟状幼体和幼水母的生长、发育、变态及成活的影响进行实验研究,发现浮浪幼虫生存的盐度下限为12‰;螅状幼体生存的盐度下限为10‰;最适盐度范围为14—20‰;碟状幼体生存的盐度下限为8‰,最适盐度范围为14—20‰;幼水母生存的盐度下限为8‰。 作者根据近15年辽东湾北部主要河流的淡水注入量,海蜇产量及本实验数据,结合辽东湾曾出现过三次大减产现象分析,认为前一年夏、秋季海蜇生殖季节河流淡水注入量过多,使河口附近海蜇浮浪幼虫和螅状幼体的栖息水域盐度聚降,导致浮浪幼虫和螅状幼体毁灭性死亡,是造成海蜇资源锐减的主要原因。作者并提出采取人工培育螅状幼体,选择合适水域进行放流以补充海蜇资源,是海蜇渔业稳定和增产的主要途径。     

大型水母幼体生长的影响因子研究进展

21世纪以来,中国东、黄海,韩国西海岸以及日本海连年发生大型水母暴发现象,对海洋渔业的生产活动以及海洋生态系统带来巨大的影响。水母暴发形成机制非常复杂,解释其发生机理并有效预报是目前急待解决的问题。大型水母的生活史中有明显的世代交替现象,受精卵,浮浪幼虫,螅状体,足囊,横裂体到碟状体的幼体发育阶段属无性世代,幼蜇发育到成蜇阶段属有性世代。在早期生活史中,螅状体的足囊繁殖与横裂生殖是大型水母无性繁殖的重要方式,对其成体的数量形成至关重要。综述了国内外有关温度、盐度、光以及营养条件对大型水母早期发育阶段的影响研究进展,研究表明温度是影响螅状体发育以及足囊繁殖和横裂生殖的最主要的环境因子;盐度、光和营养条件在适温范围内,均对螅状体和横裂生殖有一定的影响,其上下限随水母种类和发育阶段有所变化。展望了大型水母早期幼体研究的发展趋势,如环境因子对不同种类的大型水母幼体生长机理的影响、多个环境因子对幼体的综合作用、动态的环境因子与大型水母幼体之间的关系等。     

温度和食物水平对海月水母螅状体无性繁殖的影响

水母暴发给近岸人类的生活、渔业资源以及海洋生态系统带来影响。这些近岸海域暴发的水母可以通过有性繁殖和无性繁殖来维持或扩大水母种群数量。在水母生活史中,螅状体的无性繁殖是决定水母体数量的关键阶段,因此对此阶段进行研究。设置了4个温度水平(9、12、15、18℃)、3个食物水平(5个卤虫/螅状体、20个卤虫/螅状体、40个卤虫/螅状体),在12个组合条件下研究温度和食物水平对海月水母螅状体无性繁殖能力和方式的影响。研究结果表明,在海月水母螅状体繁殖子体的各种方式中,匍匐茎生殖是主要的繁殖方式,出芽生殖次之,纵向分裂以及足囊出现几率极低。食物对海月水母螅状体产生总子体数影响显著,温度的影响不显著,食物水平越高,海月水母螅状体繁殖子体的能力越强。食物和温度对螅状体发生横裂均有影响,但温度对螅状体横裂的影响更大。温度对螅状体的横裂率影响显著,食物影响不显著。碟状体的释放发生在12、15、18℃的条件下,温度是影响海月水母螅状体通过横裂生殖释放碟状体数量的最重要因素。可见在螅状体无性繁殖阶段,温度和食物对繁殖方式的影响各不相同。     

环境因子对海月水母生长发育影响的研究进展

海月水母是一种广泛分布的世界性近岸种,也是我国近海海域大型水母优势种之一.近年来海月水母在世界各地频繁暴发,对海洋生态系统和沿岸经济、社会发展带来了很大危害.海月水母有复杂的生活史(营附着生活的螅状体世代和营浮游生活的水母体世代)和多种无性生殖(出芽生殖、横裂生殖、足囊生殖等)及有性生殖方式.环境因子影响海月水母生活史的各个阶段,如浮浪幼虫阶段、螅状体阶段、碟状体阶段等.这些生长阶段尤其是早期生长阶段的状态(如附着、分布、扩散等)会进一步影响海月水母的种群动态.本文以海月水母生活史为主线,系统总结分析了国内外关于环境因子对其生活史各阶段的影响,提出了一些值得进一步深入研究的问题,以期为我国近海海域水母暴发关键因子的研究提供借鉴.     

温度、投饵频次对白色霞水母无性繁殖与螅状体生长的影响

白色霞水母是我国近海主要大型灾害水母种类之一,其暴发性增殖严重破坏了海洋生态系统平衡.在室内控制条件下,研究了温度(7.5、11、14.5、18、21.5和25℃)和投饵频次(1次/2d、1次/8d和1次/16d)对白色霞水母无性繁殖与螅状体生长的影响.结果显示,白色霞水母足囊繁殖的适宜温度为18-25℃,足囊繁殖随温度和投饵频次的增加而增加.温度对白色霞水母横裂率和横裂次数的影响显著,温度越高,白色霞水母发生横裂生殖的时间越早,横裂生殖速度越快,重复横裂次数越多,释放的碟状体数量也越多.横裂率和横裂次数随投饵频次的增加而递增.白色霞水母螅状体在7.5-25℃范围的成活率均为100％,其生长速度随温度和投饵频次的增加而增加.温度和投饵频次对白色霞水母螅状体足囊繁殖、横裂率和螅状体生长具有明显的交互效应.螅状体的横裂次数和初生碟状幼体伞径随螅状体柄径增大而递增,呈线性相关.研究表明,温度、投饵频次即营养条件显著影响着白色霞水母的种群数量,说明海水水温上升、富营养化或渔业资源锐减导致的浮游动物量增加均可能诱发白色霞水母暴发性增殖.结论为进一步探索大型水母暴发的生态环境机理提供重要科学依据.     

盐度对不同温度下海月水母螅状体存活和无性繁殖的影响

海月水母是全球近岸海域的主要致灾水母种类之一,其螅状体的繁殖情况与种群数量是影响水母暴发的重要因素.采用实验生态学的方法分别研究了高温(21 ℃)和低温(12 ℃)条件下,不同盐度梯度对螅状体存活与无性繁殖的影响.结果表明: 高温条件下,盐度15～40时螅状体存活率均大于90%,适合出芽生殖盐度范围为20～32,其中28为最适盐度;在低盐(≤15)或高盐(≥36)环境下,螅状体会进行足囊生殖以度过不良环境条件. 低温条件下,20～40盐度组螅状体存活率均大于90%,20～32盐度组适于螅状体出芽生殖,其中28盐度组出芽生殖效率最高;20～40盐度适于横裂生殖,其中28～32盐度组最利于螅状体横裂生殖. 说明海月水母螅状体有较强的盐度耐受性,一定范围内盐度对螅状体无性生殖影响不显著.     

FMRF-amide immunoreactivity pattern in the planula and colony of the hydroid Gonothyraea loveni

Gonothyraea loveni (Allman, 1859) is a colonial thecate hydrozoan with a life cycle that lacks a free-swimming medusa stage. The development from zygote to planula occurs within meconidia attached to the female colony. The planula metamorphosis results in the formation of a primary hydranth. The colony then grows by development of new colony elements. In the present work, we studied the temporal pattern of the formation of FMRF-amide-positive cells during embryogenesis, in larvae and during early colony ontogeny. FMRF-amide-positive cells appear in the planula only after its maturation. However, they disappear after planula settlement. For the first time, we show that neural cells are present in the coenosarc of the hydroid colony. We also trace the process of neural net formation during the development of a new shoot internode of the G. loveni colony.     

The effect of larval age on developmental changes in the polyp prepattern of a hydrozoan planula

The larvae of many marine organisms including hydrozoans are lecithotrophic and will not feed until after metamorphosis. In hydrozoans the aboral region of the planula becomes the holdfast and stolon, while the oral region becomes the stalk and hydranth that grows out of the holdfast following metamorphosis. If metamorphosis is delayed, the portion of the planula allocated to form holdfast and stolon shrinks and the region that forms the hydranth increases in size. Planulae also have the ability to regenerate their polyp prepattern. When the aboral region of the planula that does not normally form a hydranth is isolated and metamorphosis is delayed, it acquires the capacity to form a hydranth from the holdfast. A relatively high proportion of entodermal cells of young planulae engage in DNA synthesis (BrdU labeling index); as planulae age, the labeling index falls close to zero. When the polyp prepattern is modified during planula regeneration, entodermal cells are induced to engage in DNA synthesis. If DNA synthesis is inhibited in planulae, the polyp prepattern changes during regeneration and age-related developmental changes in planula are inhibited, suggesting that DNA synthesis is a necessary part of the pattern respecification process.     

Experimentelle Untersuchungen zum Stockwachstum und zur Medusenbildung bei dem marinen HydrozoonEirene viridula

The development of both slide-grown and non-substrate bound colonies ofE. viridula (Thecata-Leptomedusae) ranging in size from 1 to 50 hydranths was investigated under various temperature conditions. The majority of slide-grown colonies reached a larger final size than non-substrate bound ones, in 20°, 25° and 29° C. Raising the temperature did not stimulate propagation of hydranths as expected. Most of the colonies transferred to 25° or 29° C finally were even smaller than those reared at 20° C. This was partially due to resorption of several hydranths about 9 days after the temperature rise; the influence of “physiological competition” between development of new hydranths and budding of medusae on colony growth is discussed. Transfer from higher to lower temperatures affected colony growth negatively. Raising the temperature from 20° to 25° or 29° C initiated formation of gonozooids from the distal part of hydranth stalks and development of medusa buds in both types of colonies. With the exception of slide-grown colonies transferred to 25° C, also young medusae were budded off. There was a remarkable coincidence in predominance of colony growth in slide-grown colonies and of medusa budding in non-substrate bound cultures. In the latter, medusa buds developed 1 to 2 days earlier. Most buds did not differentiate into liberated medusae, but were resorbed. Transformation of medusa buds into hydranths was not observed. In the clone ofE. viridula, onset of medusa budding did not depend on a “minimal colony size”. Even single hydranths were able to produce medusa buds after transfer to higher temperatures; budded off medusae were recorded from non-substrate bound colonies with an initial size of 3 hydranths. In slide-grown cultures, medusa buds developed into colonies with an initial size of only 3 hydranths. No hydranth propagation prior to medusa budding occurred in these cases. After raising temperature from 25° to 29° C medusa buds were observed in nonsubstrate bound colonies only; a small number of medusae were budded off from some of these colonies. Lowering the temperature from 29° or 25° to 20° C caused resorption of existing medusa buds. In several non-substrate bound colonies, transfer from 29° to 25° C induced development of gonozooids with medusa buds and, in some cases, of young medusae. Incubation with the alkylating cytostaticTrenimon and transfer from 20° to 25° C caused irreversible resorption of all hydranths when 4 × 10^?2 mg/ml were administered for 10 mins. Thereafter, only development of stolonial structures was observed. With one exception, the colonies treated with 4 × 10^?3 mg/ml, and all others submitted to 4 × 10^?4 mgTrenimon/ml were able to produce new hydranths and also medusa buds; some of the colonies first had to overcome a degressive phase. Treatment with 4 × 10^?2 mg destroyed all interstitial cells (I-cells). Incubation with 1 × 10^?3 or 1 × 10^?4 mg/ml left the I-cells at least partially intact. It is concluded that I-cells are indispensable for hydranth and medusan morphogenesis inE. viridula.     

Is the remodeling of the polyp prepattern in a hydrozoan planula a function of larval age or size and is it of “adaptive” significance?

Eggs of medusae develop into lecithotrophic planulae that undergo metamorphosis at different ages to form polyps. As planulae age they decrease in size as their yolk stores are utilized. The planulae of most Phialidium medusae develop into polyps where there is a decrease in the size of the holdfast region and a relative increase in the size of the hydranth region as they age. These changes occur independently of the decrease in planula size. In planulae with a decrease in the size of the holdfast region and an increase in the size of the hydranth-forming region there was a 50% decline in polyps that successfully stayed attached to the substrate after metamorphosis. These aged planulae produced an initial hydranth with the same number of tentacles as polyps from full-sized young planulae while young half-sized planulae produced hydranths where the tentacle number was smaller. The first phase of polyp colony growth with a small initial hydranth was slower than growth of a colony with a larger initial hydranth. Predation during this period led to more death in colonies with a small initial hydranth. The decline in successful attachment in aged planulae was not offset by the higher rate of growth and a smaller window of time where predation leads to death, suggesting that this age-related developmental change in planulae was not adaptive.     

Relay-ray way of hydroplasm movement in hydroid colonies

In this study, we continued the investigation of the distribution system of colonial hydroids in the course of its development, starting with its emergence during the planula metamorphosis and ending with the formed colony. The hydroplasmic stream system of two species of colonial hydroids—Perigonimus abyssi G.O. Sars, 1874, and Stauridia producta Wright, 1858—was studied. We found that the main principle by which hydroplasma moves in these colonies, which form no shoots, is the relay-race from one hydranth to another through a stolon fragment connecting them or it moves past the neighbouring hydranth to the next one, etc. We show that the efficiency of the conducting (distribution) system does not depend on the level of complexity of the colonial structure. The results of this study confirm the absence of general colonial processes of integration and self-regulation in hydroid polyps.     

Reproduction of the colonial hydroid Obelia geniculata (L., 1758) (Cnidaria,Hydrozoa) in the White Sea

Populations of the colonial hydroid Obelia geniculata in the White Sea reproduce asexually by frustule formation. Young medusae appear in the plankton during July and August. The number of medusae rarely exceeds 36 per m³, and the average number varies every year from 0.4 to 10 per m³. The size of medusae is smaller than reported from other regions. The umbrella of the largest recorded medusa was only 0.57 mm in diameter and the specimen had just 35 tentacles. Only a few mature medusae were found during the study. The colonies in the White Sea are epiphytic and grow only on laminarian thalli. At the beginning of July there are no colonies on thalli from the upper subtidal zone. By the end of August, colonies of O.␣geniculata had increased in density to 30 per m². Hydroid recruitment was attributed to active frustule production by colonies living below that zone. The frustules detach from the stems of the hydroids and are found in plankton. Production of frustules on branches occurs continuously during colony growth until water temperatures climb above 0 °C. We found that water temperature in this Arctic environment is generally too low for medusa maturation and planula development in the species. Propagation by frustule formation is the principal means of reproduction in Obelia geniculata within the White Sea, and this phenomenon accounts for the species being a dominant epiphyte on laminarian thalli there.     

Nematocysts and Taxonomy in Laomedea, Gonothyraea and Obelia (Hydrozoa, Campanulariidae)

The cnidoms of Laomedea flexuosa, Gonothyraea loveni, Obelia geniculata, O. longissima and O. dichotoma were studied by light and scanning electron microscopy to find out whether the nematocysts could be used as taxonomic characters. Three b-rhabdoid heteroneme nematocysts (microbasic b-mastigophore) and three isorhizous haploneme nematocysts (atrichous isorhiza) were distinguished. A small b-rhabdoid nematocyst with spindle-shaped capsule occurred in all the species examined. In the polyp and planula of G. loveni , and the planula of L. flexuosa it was the only nematocyst present. Specific for L. flexuosa was a b-rhabdoid with curved capsule. In the polyp and newly-liberated medusa of O. longissima another b-rhabdoid appeared with bean-shaped capsule and markedly long spines at the distal tube. The polyp and newly-liberated medusa of O. dichotoma were characterized by two different isorhizas. A special type of isorhiza occurred in the polyp of O. geniculata . The curved capsules of the different isorhizas varied somewhat in shape and size. Differences in nematocyst structure and occurrence are presumed to provide characters for taxonomy. Thus, O. dichotoma and O. Iongissima are regarded as separate species due to their distinctly different nematocysts, in conjunction with other morphological and ecological differences.     

Phylogenetic analysis of the Cnidaria

The recent members of the phylum Cnidaria were analyzed with phylogenetic methodology and the help of the PAUP Computer program. The Cnidaria are established as a monophylum by their cnidocysts, planula larva, and a polyp stage. The Ctenophora were seen as the most probable sister group of the Cnidaria. Arguments for the monophyly of the cnidarian classes Anthozoa, Scyphozoa, Cubozoa, and Hydrozoa were providea. For the ground plan of the Cnidaria the following characters were postulated: triphasic life cycle consisting of a planula larva, a benthic polyp stage, and a sexually propagating medusa like stage. For the polyp a radial symmetry, lack of septae, and hollow tentacles were assumed. The original medusa probably was tetraradial and developed from the polyp stage by a total metamorphosis. Twelve polarized characters were used to generate cladograms. The most parsimonious one showed the Anthozoa as the first offshoot of the tree with the united Scyphozoa, Cubozoa and Hydrozoa forming its sister group. Within this sister group the Scyphozoa and Cubozoa were seen as sistergroups to each other. Both groups united are then the sistergroup of the Hydrozoa. A bootstrap analysis yielded the same tree with high probabilities for the internal nodes. Despite assuming a planktonic origin of the Cnidaria in this investigation, the resulting cladogram is also compatible with an evolution of the medusa stage within the Cnidaria after the splitting-off of the Anthozoa. The possible loss of the medusa stage in the Anthozoa is discussed.