鼠尾藻（Sargassum Thunbergii）繁殖生态学研究
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摘要
鼠尾藻种群更新取决于三个生活史阶段：生殖分配、配子生殖和幼殖体补充，本文以生活史为线索，围绕这三个阶段开展了繁殖生态学研究。
烟台地区鼠尾藻一年一次的有性生殖时期自6月中旬开始，7月中旬达高峰，此时90％藻体成熟，生殖分配(RA)均值达到75％。在繁殖季节，生殖分配和成熟比例均显示了显著的时间变化。繁殖高峰时，未成熟的藻体仅出现在小长度级，生殖分配均值显示了显著的层级变化，two-wayANOVA分析结果表明发育进程和藻体长度对藻体成熟和生殖分配具有显著影响，证实了成熟和生殖分配的大小依赖。侧枝的产生具有明显的季节性，并在侧枝产生后有性生殖启动。生殖分配与侧枝数量、长度呈正相关，意味着侧枝的出现对生殖具有重要意义。另外，有性生殖季节过后，大量的构件(module)开始脱落，而且成熟藻体构件的脱落明显多于未成熟的藻体。因为在7-8月所有存活下来的藻体都是未成熟的小个体。所以，生殖和生长的权衡关系(trade-off)在鼠尾藻个体水平上可能是存在的。
海藻配子释放得到了很多的关注，但引起配子释放的环境因子间的交互作用以及各种因子对配子释放的相对重要性却没有得到足够的重视。本文采用了2~(8-4)分式析因设计(ResolutionⅣ)侦寻环境因素的主效应和交互作用。结果显示，在设计的因素中，水流、温度、盐度和光期是影响配子释放的重要因素，这些因素间的两维交互作用均显著。基于方差分析评估的影响效应，环境因素对配子释放的相对重要性依次为盐度、水流*光期(或温度*盐度)、水流*盐度(或温度*光期)、温度、水流*温度(或光期*盐度)、水流和光期。最适的因素组合25℃、30‰盐度、静水、12 h光照可能是鼠尾藻配子释放的“机会窗”。这些实验室条件下的研究结果对于深入认识配子释放机制具有重要意义。
野外跟踪观测鼠尾藻的成熟繁殖状况，采集临近成熟的鼠尾藻至实验室培养，跟踪观测其卵子和受精卵的发育以及幼殖体的早期生长。研究发现，鼠尾藻的卵子发育属于8核1卵型，卵子受精后20 h左右开始脱离生殖托，受精卵先连续进行两次的横分裂，而后再由一端的一个细胞发育成假根，另外两个发育成体细胞。幼孢子体在接近2mm时候开始形成分枝突起。
同步的配子释放和杂藻的生长是鼠尾藻苗种人工繁殖中的两个难题。自然群体和人工养殖群体比较研究结果表明，在繁殖高峰期自然群体和人工养殖群体生殖力分别为40％和65％；幼殖体发生呈现显著的时间变化，不过，相对自然群体，养殖群体显示了较低的时间离散。平均每公斤鲜重的养殖群体获得了约3．2×10~5株幼殖体，可以满足大规模生产的需要。本文采用了高密度播种和洗刷的办法以消除杂藻的生长。幼殖体播种后的附着能力随时间逐渐提高。播种后48h采用1 kg cm~(-1)的压力洗刷育苗器时，脱落率不足10％，使洗刷可以在生活史早期阶段进行，从而在早期阶段就开始阻止杂藻的生长。研究发现，高密度的播种对幼殖体生长具有消极的影响：减小了平均长度、大小不均性提高、侧枝发生偏晚。然而，30-50 cm~(-2)的播种密度下，幼殖体受到的种内竞争的影响较小，并且可以忍受连续的洗刷。在结合高密度和洗刷的处理中，杂藻被有效消除，表明了这两项措施是有效的。经过1个月的培养多数幼殖体长度达到0．5 cm以上。
潮间带生态学调查了鼠尾藻种群结构的季节变化，采用逐步回归分析了生境因子对生物量的影响。结果表明，鼠尾藻分布在低潮线以上50～125cm的中潮带和低潮带之间；生物量消长和平均藻体长度消长的季节变化趋势一致，呈双峰曲线(7月份和12月份达到峰值，9月份达到最低值)；生活周期可以划分为4个时期：休止期、生长期、繁殖期和衰退期，6月中旬到7月下旬为有性生殖期；营养生殖贯穿全年，并呈现一定的季节变化；快速生长期内(5～7月)，其大小级层次明显，近似正态分布；水温为影响其生长的主要因素，其次为浪冲击度和人为干扰，干露对鼠尾藻的生长影响不显著。总之，本调查区域的鼠尾藻种群呈现明显的季节变化，并与其它地区种群差异显著，生长地区的环境因素是导致差异的根本原因。
Three life-history stages including reproductive allocation (RA),gametogenesisand recruitment influence the population structure of Sargassum thunbergii.Based onthis life-history cycle,the reproductive ecology of the species was conducted.
Annual reproduction initiated in mid-June and peaked in mid-July (90% fertilethalli and 75% RA mean).Both RA and percentage of fertile thalli exhibitedsignificant temporal variations during reproduction.Sterile thalli were only observedin small hierarchies at peak reproduction and mean values of RA showed asignificantly hierarchical variation,suggesting that the size of thalli played importantroles in reproduction and both reproduction and RA were size-dependent.Numerousreceptacles were produced along the lateral branches during the reproductive period.A distinct seasonal pattern was observed wherein the presence of lateral branches wasfollowed by the onset of reproduction.RA was positively correlated with the numberof lateral branches,as well as the total length of lateral branches,suggesting that theappearance of lateral branches is of importance for reproduction.In addition,massiveshedding of modules occurred after the reproductive season.Moreover,theprobability of shedding was evidently higher for fertile than for sterile modules,because all surviving thalli were sterile and short during July-August.So,a trade-offbetween reproduction and survival may exist at individual levels in S.thunbergii.
More attentions have been paid to the gamete release of fucoid algae,but theinteractions between the environmental cues and their importance order for thegamete release remain little known.In this study,a 2~(8-4) fractional factorial design(ResolutionⅣ) was used to identify the main effects and interactions ofenvironmental factors.Of the factors tested,water motion,temperature,photoperiodand salinity were found to be the most important factors affecting egg release.Moreover,the interactions between any two factors were all significant.Based on thesize of effect estimated by ANOVA,the effects which contribute to the release ofeggs,in order of importance,are:salinity,water motion~*photoperiod (possiblytemperature~*salinity),water motion~*salinity (possibly temperature~*photoperiod),temperature,water motion~*temperature (photoperiod~*salinity),water motion and photoperiod.The optimum conditions of 25℃,30‰salinity,slack seawater and 12 hirradiance that achieved 100% release rate may be the“window of opportunity”ofegg release in S.thunbergii.These findings give insight into the egg release in S.thunbergii under laboratory conditions.
The development of ovum and zygote,and the growth of juvenile sporophyte wereobserved on the laboratory conditions.Eggs adhered to the receptacle followingrelease from their conceptacles.The zygote detached from receptacle 20 happroximately after fertilization.After taking transverse division twice,one terminalcell developed into rhizoid and the two others developed into the germling.Thejuveniles started growing new branch when nearly reached 2mm in length.
Commercial farming of Sargassum thunbergii is being developed in China.Asynchronous discharge of gametes and growth of epiphytic algae are the two mainconstraints for the seedling production of this species.In this study,40% and 65%reproductive output for nature and farmed population were respectively recordedduring peak reproduction.Furthermore,lower degree of temporal dispersion ofgermling occurrences for farmed population in related to nature ones was found,although germling occurrence exhibited significant temporal variations for bothpopulations.These results indicated that using farmed population as parental plants tocollect germlings is an effective strategy in achieving mass discharge of gametes.Atthe peak period of discharge,as more as 3.2×10~5 germlings per kg wet weight plantsfrom the farmed population was obtained,which was significantly more than that ofnature ones and could meet the demand of large scale seedling production.Anotherconstraint (i.e.growth of epiphytic algae) interfering the seedling cultivation wasattempted to eliminate by a strategy of jet-washing and high density seeding.Theattachment capacity of germlings was found to be significantly increased with theduration of attachment.A detachment of less than 10% was observed on collectorsthat were jet-washed using an intensity of 1 kg cm~(-1) by 48 h post-attachment,whichenables the jet-washing to be early conducted,thus preventing the growth of epiphyticalgae from early life stage.High density had adverse effects on length mean,sizeequality and occurrence of branches of germlings.However,30-50 individuals cmwere thought to be usable in the seedling production of S.thunbergii,because theyunderwent less density effects,and withstood the impact of continual jet washing.Noevident epiphytic algae growing on collectors or adhering to host plants were observed in the treatments which combined jet-washing and high seeding density,suggesting that both jet-washing and high density significantly reduced epiphyticalgae.Most seedlings in a length of＞0.5 cm were achieved after one month of tankculture under less controlled conditions.
The effects of environmental factors including disturbance,temperature,wavemotion,epiphytes and grazer (i.e.Chlorostorna rustica) on biomass were conducted,using step-wise regression analysis.The results showed that S.thunbergii distributedbetween the low-tidal zone and mid-tidal zone at 50～125 cm tidal level.Both biomassand mean thallus length of S.thunbergii exhibited a statistically significant temporalvariation,with a similar unimodal trajectory.The biomass and mean thallus lengthreached a highest value in July and decreased to a lowest value in September.Thegrowth pattern of S.thunbergii may be divided into four phases:inactivity period(before March),growth period (from early-April to mid-June),reproductive period(from mid-June to late-July) and senescence period (from late-July to September).Small plants were recorded throughout the year and showed a seasonal variation,indicating that the vegetative reproduction ofS.thunbergii occurred year round.AfterJuly 2005,the number of smallest plants increased rapidly due to the degeneration oflarge plants,recruitment of new plants and vegetative reproduction.During therapid-growth period,the length distribution was not dominated by a few size classes,and-frequencies were more or less evenly distributed in most size classes.Step-wiseregression analysis demonstrated that water temperature,wave motion anddisturbance significantly influenced the increase of biomass;however,the effect ofexposure was not significant.That is considered to be the results that S.thunbergiiclumped during sampling period,because fronds covered each other at low-tide andwater lose was efficiently prevented.On balance,the population investigated inpresent study is significantly different from others growing on the coast of Japan andKorea,which results from the differences of environmental factors.
烟台地区鼠尾藻一年一次的有性生殖时期自6月中旬开始，7月中旬达高峰，此时90％藻体成熟，生殖分配(RA)均值达到75％。在繁殖季节，生殖分配和成熟比例均显示了显著的时间变化。繁殖高峰时，未成熟的藻体仅出现在小长度级，生殖分配均值显示了显著的层级变化，two-wayANOVA分析结果表明发育进程和藻体长度对藻体成熟和生殖分配具有显著影响，证实了成熟和生殖分配的大小依赖。侧枝的产生具有明显的季节性，并在侧枝产生后有性生殖启动。生殖分配与侧枝数量、长度呈正相关，意味着侧枝的出现对生殖具有重要意义。另外，有性生殖季节过后，大量的构件(module)开始脱落，而且成熟藻体构件的脱落明显多于未成熟的藻体。因为在7-8月所有存活下来的藻体都是未成熟的小个体。所以，生殖和生长的权衡关系(trade-off)在鼠尾藻个体水平上可能是存在的。
海藻配子释放得到了很多的关注，但引起配子释放的环境因子间的交互作用以及各种因子对配子释放的相对重要性却没有得到足够的重视。本文采用了2~(8-4)分式析因设计(ResolutionⅣ)侦寻环境因素的主效应和交互作用。结果显示，在设计的因素中，水流、温度、盐度和光期是影响配子释放的重要因素，这些因素间的两维交互作用均显著。基于方差分析评估的影响效应，环境因素对配子释放的相对重要性依次为盐度、水流*光期(或温度*盐度)、水流*盐度(或温度*光期)、温度、水流*温度(或光期*盐度)、水流和光期。最适的因素组合25℃、30‰盐度、静水、12 h光照可能是鼠尾藻配子释放的“机会窗”。这些实验室条件下的研究结果对于深入认识配子释放机制具有重要意义。
野外跟踪观测鼠尾藻的成熟繁殖状况，采集临近成熟的鼠尾藻至实验室培养，跟踪观测其卵子和受精卵的发育以及幼殖体的早期生长。研究发现，鼠尾藻的卵子发育属于8核1卵型，卵子受精后20 h左右开始脱离生殖托，受精卵先连续进行两次的横分裂，而后再由一端的一个细胞发育成假根，另外两个发育成体细胞。幼孢子体在接近2mm时候开始形成分枝突起。
同步的配子释放和杂藻的生长是鼠尾藻苗种人工繁殖中的两个难题。自然群体和人工养殖群体比较研究结果表明，在繁殖高峰期自然群体和人工养殖群体生殖力分别为40％和65％；幼殖体发生呈现显著的时间变化，不过，相对自然群体，养殖群体显示了较低的时间离散。平均每公斤鲜重的养殖群体获得了约3．2×10~5株幼殖体，可以满足大规模生产的需要。本文采用了高密度播种和洗刷的办法以消除杂藻的生长。幼殖体播种后的附着能力随时间逐渐提高。播种后48h采用1 kg cm~(-1)的压力洗刷育苗器时，脱落率不足10％，使洗刷可以在生活史早期阶段进行，从而在早期阶段就开始阻止杂藻的生长。研究发现，高密度的播种对幼殖体生长具有消极的影响：减小了平均长度、大小不均性提高、侧枝发生偏晚。然而，30-50 cm~(-2)的播种密度下，幼殖体受到的种内竞争的影响较小，并且可以忍受连续的洗刷。在结合高密度和洗刷的处理中，杂藻被有效消除，表明了这两项措施是有效的。经过1个月的培养多数幼殖体长度达到0．5 cm以上。
潮间带生态学调查了鼠尾藻种群结构的季节变化，采用逐步回归分析了生境因子对生物量的影响。结果表明，鼠尾藻分布在低潮线以上50～125cm的中潮带和低潮带之间；生物量消长和平均藻体长度消长的季节变化趋势一致，呈双峰曲线(7月份和12月份达到峰值，9月份达到最低值)；生活周期可以划分为4个时期：休止期、生长期、繁殖期和衰退期，6月中旬到7月下旬为有性生殖期；营养生殖贯穿全年，并呈现一定的季节变化；快速生长期内(5～7月)，其大小级层次明显，近似正态分布；水温为影响其生长的主要因素，其次为浪冲击度和人为干扰，干露对鼠尾藻的生长影响不显著。总之，本调查区域的鼠尾藻种群呈现明显的季节变化，并与其它地区种群差异显著，生长地区的环境因素是导致差异的根本原因。
Three life-history stages including reproductive allocation (RA),gametogenesisand recruitment influence the population structure of Sargassum thunbergii.Based onthis life-history cycle,the reproductive ecology of the species was conducted.
Annual reproduction initiated in mid-June and peaked in mid-July (90% fertilethalli and 75% RA mean).Both RA and percentage of fertile thalli exhibitedsignificant temporal variations during reproduction.Sterile thalli were only observedin small hierarchies at peak reproduction and mean values of RA showed asignificantly hierarchical variation,suggesting that the size of thalli played importantroles in reproduction and both reproduction and RA were size-dependent.Numerousreceptacles were produced along the lateral branches during the reproductive period.A distinct seasonal pattern was observed wherein the presence of lateral branches wasfollowed by the onset of reproduction.RA was positively correlated with the numberof lateral branches,as well as the total length of lateral branches,suggesting that theappearance of lateral branches is of importance for reproduction.In addition,massiveshedding of modules occurred after the reproductive season.Moreover,theprobability of shedding was evidently higher for fertile than for sterile modules,because all surviving thalli were sterile and short during July-August.So,a trade-offbetween reproduction and survival may exist at individual levels in S.thunbergii.
More attentions have been paid to the gamete release of fucoid algae,but theinteractions between the environmental cues and their importance order for thegamete release remain little known.In this study,a 2~(8-4) fractional factorial design(ResolutionⅣ) was used to identify the main effects and interactions ofenvironmental factors.Of the factors tested,water motion,temperature,photoperiodand salinity were found to be the most important factors affecting egg release.Moreover,the interactions between any two factors were all significant.Based on thesize of effect estimated by ANOVA,the effects which contribute to the release ofeggs,in order of importance,are:salinity,water motion~*photoperiod (possiblytemperature~*salinity),water motion~*salinity (possibly temperature~*photoperiod),temperature,water motion~*temperature (photoperiod~*salinity),water motion and photoperiod.The optimum conditions of 25℃,30‰salinity,slack seawater and 12 hirradiance that achieved 100% release rate may be the“window of opportunity”ofegg release in S.thunbergii.These findings give insight into the egg release in S.thunbergii under laboratory conditions.
The development of ovum and zygote,and the growth of juvenile sporophyte wereobserved on the laboratory conditions.Eggs adhered to the receptacle followingrelease from their conceptacles.The zygote detached from receptacle 20 happroximately after fertilization.After taking transverse division twice,one terminalcell developed into rhizoid and the two others developed into the germling.Thejuveniles started growing new branch when nearly reached 2mm in length.
Commercial farming of Sargassum thunbergii is being developed in China.Asynchronous discharge of gametes and growth of epiphytic algae are the two mainconstraints for the seedling production of this species.In this study,40% and 65%reproductive output for nature and farmed population were respectively recordedduring peak reproduction.Furthermore,lower degree of temporal dispersion ofgermling occurrences for farmed population in related to nature ones was found,although germling occurrence exhibited significant temporal variations for bothpopulations.These results indicated that using farmed population as parental plants tocollect germlings is an effective strategy in achieving mass discharge of gametes.Atthe peak period of discharge,as more as 3.2×10~5 germlings per kg wet weight plantsfrom the farmed population was obtained,which was significantly more than that ofnature ones and could meet the demand of large scale seedling production.Anotherconstraint (i.e.growth of epiphytic algae) interfering the seedling cultivation wasattempted to eliminate by a strategy of jet-washing and high density seeding.Theattachment capacity of germlings was found to be significantly increased with theduration of attachment.A detachment of less than 10% was observed on collectorsthat were jet-washed using an intensity of 1 kg cm~(-1) by 48 h post-attachment,whichenables the jet-washing to be early conducted,thus preventing the growth of epiphyticalgae from early life stage.High density had adverse effects on length mean,sizeequality and occurrence of branches of germlings.However,30-50 individuals cmwere thought to be usable in the seedling production of S.thunbergii,because theyunderwent less density effects,and withstood the impact of continual jet washing.Noevident epiphytic algae growing on collectors or adhering to host plants were observed in the treatments which combined jet-washing and high seeding density,suggesting that both jet-washing and high density significantly reduced epiphyticalgae.Most seedlings in a length of＞0.5 cm were achieved after one month of tankculture under less controlled conditions.
The effects of environmental factors including disturbance,temperature,wavemotion,epiphytes and grazer (i.e.Chlorostorna rustica) on biomass were conducted,using step-wise regression analysis.The results showed that S.thunbergii distributedbetween the low-tidal zone and mid-tidal zone at 50～125 cm tidal level.Both biomassand mean thallus length of S.thunbergii exhibited a statistically significant temporalvariation,with a similar unimodal trajectory.The biomass and mean thallus lengthreached a highest value in July and decreased to a lowest value in September.Thegrowth pattern of S.thunbergii may be divided into four phases:inactivity period(before March),growth period (from early-April to mid-June),reproductive period(from mid-June to late-July) and senescence period (from late-July to September).Small plants were recorded throughout the year and showed a seasonal variation,indicating that the vegetative reproduction ofS.thunbergii occurred year round.AfterJuly 2005,the number of smallest plants increased rapidly due to the degeneration oflarge plants,recruitment of new plants and vegetative reproduction.During therapid-growth period,the length distribution was not dominated by a few size classes,and-frequencies were more or less evenly distributed in most size classes.Step-wiseregression analysis demonstrated that water temperature,wave motion anddisturbance significantly influenced the increase of biomass;however,the effect ofexposure was not significant.That is considered to be the results that S.thunbergiiclumped during sampling period,because fronds covered each other at low-tide andwater lose was efficiently prevented.On balance,the population investigated inpresent study is significantly different from others growing on the coast of Japan andKorea,which results from the differences of environmental factors.
引文
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