轮虫同域性物种形成:来自萼花臂尾轮虫克隆间的分子系统发育关系和生殖隔离的证据

对采自芜湖市三个不同水体中的萼花臂尾轮虫(Brachionus calyciflorus)17个克隆的线粒体COⅠ基因和rDNAITS1序列进行的分析结果表明:萼花臂尾轮虫不同克隆间COⅠ基因序列差异百分比为0-14.0%,平均为5.1%;ITS1序列差异百分比为0-7.4%,平均为2.2%。基于COⅠ基因序列构建的分子系统树(NJ树、MP树、ML树和贝叶斯树)均支持将17个克隆划分为三个姐妹种,各姐妹种间的序列差异百分比为7.2%-14.0%。基于ITS1序列构建的同样四种分子系统树也支持将17个克隆划分为三个姐妹种,但其中的克隆HE5在姐妹种中的归属与基于COⅠ基因序列构建的分子系统树所得出的结果不同,各姐妹种间的序列差异百分比为1.9%-7.4%。混交雌体和雄体间的交配实验结果表明,轮虫各姐妹种间存在着明显的生殖隔离,克隆HE5在姐妹种中的归属应与基于COⅠ基因序列构建的分子系统树所得出的结果一致;ITS1序列进化速率较低,不宜用于轮虫姐妹种的准确甄别。COⅠ基因进化钟计算结果显示,三个姐妹种间,克隆LE9所属的姐妹种早在8百万年前便分化形成出来,HE1、HE2、HE4-7所属的姐妹种于5百万年前分化产生。     

基于rDNA ITS序列分析莲塘湖萼花臂尾轮虫种群遗传结构的季节变化

对采自芜湖市莲塘湖水体中的萼花臂尾轮虫(Brachionus calyciflorus)夏季种群中的30个克隆(S1-S30)和秋季种群中的20个克隆(A1-A20)的rDNA ITS序列进行了分析;以十指臂尾轮虫B.patulus为外群,构建了50个克隆的NJ、MP和ME系统发生树。结果表明:三个系统树均支持将50个克隆分为两个支系;第一支系包括29个夏季采集的单元型(单元型S9除外)和1个秋季采集的单元型A2,第二个支系包括18个秋季采集的单元型(单元型A2除外)和1个夏季采集的单元型S9;两支系间ITS序列差异为5.1%-8.8%;它们应属于两个不同的姐妹种。夏季,莲塘湖水体中的萼花臂尾轮虫种复合体内以第一支系为主构成的姐妹种在相对丰度上占绝对优势,达29/30;而秋季则相反。上述结果表明:浅水湖泊中萼花臂尾轮虫种群密度的季节变动实际上主要在于各姐妹种相对丰度的季节变化;两姐妹种的种群遗传结构季节变化符合重叠模型。     

萼花臂尾轮虫种复合体内三个姐妹种的形态差异

应用单因子方差分析和判别分析方法,分别比较了采自芜湖市3个自然水体的萼花臂尾轮虫种复合体内3个姐妹种(分别以克隆HE1和HE5、克隆HE3和SE1、克隆LE9为代表)的成体和幼体的形态差异。结果表明,克隆HE3成体的各形态参数均最小,而克隆HE1和LE9成体的各形态参数间无显著的差异;克隆HE1和HE3幼体的前中棘刺间距离和后侧棘刺长度、克隆LE9和HE1幼体的前侧棘刺间距离以及前中棘刺和侧棘刺间距离均无显著性差异,轮虫幼体的其它形态参数均以克隆LE9最长,克隆HE1次之,克隆HE3最短。对克隆HE1、HE3和LE9幼体有明显差异的形态参数所作的分布图显示,它们的分布区有很高程度的重叠。以判别分析建立的判别函数对3姐妹种成体的综合判别率中形态分析Ⅰ为47.8%,形态分析Ⅱ为55.6%;对3姐妹种幼体的综合判别率中形态分析Ⅰ和Ⅱ均为71.1%;两种形态分析对克隆HE1判别准确率均较低,分别为0-76.7%和0-74.2%;依据形态特征无法准确甄别萼花臂尾轮虫姐妹种。     

基于 rDNA ITS 1序列探讨臂尾轮属轮虫的系统发生关系

本文通过对剪形臂尾轮虫、矩形臂尾轮虫、十指臂尾轮虫、红臂尾轮虫、角突臂尾轮虫、双棘臂尾轮虫、裂足臂尾轮虫和萼花臂尾轮虫等八种臂尾轮虫rDNAITS 1序列分析,并以西氏晶囊轮虫为外群,使用PAUP和贝叶斯软件分别构建臂尾轮属轮虫系统发生树( MP树、NJ树、ML树和贝叶斯树) ,以探讨臂尾轮属的系统发生关系,并解决其中的一些分类问题。结果表明:本研究所涉及的轮虫rDNAITS 1平均序列差异百分比较高,为29 %;海水和淡水臂尾轮虫被明显分为不同的进化枝;除双棘臂尾轮虫外,在淡水臂尾轮虫中具有三对前棘刺且营附着生活的种类与前棘刺少于三对且营浮游生活的种类聚在不同支系中,这与以形态特征为主所进行的系统发生研究结果基本一致;所有的系统树均支持将十指臂尾轮虫作为一个独立的支系分离出来,裂足臂尾轮虫应归入臂尾轮属。     

萼花臂尾轮虫多态性及形态型转化

在实验室条件下研究了萼花臂尾轮虫3种形态型(双短侧棘刺、单短侧棘刺和无侧棘刺)个体之间的关系。从轮虫的培养水体中随机挑出若干3种形态型雌体的幼轮虫(龄长<24h),分别培养在正常培养液和晶囊轮虫培养液过滤液中。将实验轮虫的后代(F1代)置于200倍显微镜下检查其个体的形态型。实验结果表明,在正常培养液中,每种形态型轮虫个体都可以产生3种形态型的后代。当培养在晶囊轮虫培养液过滤液中时,这3种形态型轮虫的后代都具有2个显著增长的侧棘刺。此外,这3种形态型个体的轮虫能够共存于同一培养环境中,而且每种形态型个体的轮虫都具有3种生殖类型的雌体。由此推测萼花臂尾轮虫是一个由3种基础形态型个体组成的形态型轮虫复合体。     

萼花臂尾轮虫多态性及形态型转化

在实验室条件下研究了萼花臂尾轮虫3种形态型(双短侧棘刺、单短侧棘刺和无侧棘刺)个体之间的关系。从轮虫的培养水体中随机挑出若干3种形态型雌体的幼轮虫(龄长<24 h)，分别培养在正常培养液和晶囊轮虫培养液过滤液中。将实验轮虫的后代(F1代)置于200倍显微镜下检查其个体的形态型。实验结果表明，在正常培养液中，每种形态型轮虫个体都可以产生3种形态型的后代。当培养在晶囊轮虫培养液过滤液中时，这3种形态型轮虫的后代都具有2个显著增长的侧棘刺。此外，这3种形态型个体的轮虫能够共存于同一培养环境中，而且每种形态型个体的轮虫都具有3种生殖类型的雌体。由此推测萼花臂尾轮虫是一个由3种基础形态型个体组成的形态型轮虫复合体。     

基于线粒体COⅠ基因序列探讨臂尾轮属的系统发生和几种轮虫的分类地位

通过对萼花臂尾轮虫、角突臂尾轮虫、壶状臂尾轮虫、十指臂尾轮虫、镰形臂尾轮虫、尾突臂尾轮虫和红臂尾轮虫等7种轮虫mtDNA COⅠ序列分析,并以透明囊足轮虫为外群,使用MEGA软件构建臂尾轮属轮虫系统发生树(ME树和NJ树)以探讨臂尾轮属的系统发生和该属中几个种类的分类地位.结果表明本研究所涉及的轮虫mtDNA COⅠ平均序列差异百分比为24%;在两个系统树中,淡水臂尾轮虫和海水臂尾轮虫均独立成枝;用两种方法得到的系统发生树均支持将十指臂尾轮虫作为一个独立的支系分离出来.壶状臂尾轮虫和红臂尾轮虫应属于不同的种;来自芜湖、墨西哥和美国的十指臂尾轮虫以及来自芜湖和澳大利亚的萼花臂尾轮虫应分别互为姐妹种.本研究还首次揭示了角突臂尾轮虫、尾突臂尾轮虫和镰状臂尾轮虫与其它臂尾轮虫间的系统关系.     

剪形臂尾轮虫形态的时空变化及其与生态因子间的关系

对芜湖市镜湖和汀棠湖中剪形臂尾轮虫(Brachionus forficula)的发生及其形态特征进行了一年的观察,并分析了其与环境因子间的关系。结果表明,镜湖中,剪形臂尾轮虫出现于7月至10月,且仅存在短后棘刺型个体;2样点内,除了前外棘刺长以外,轮虫的其他各形态参数均随季节显著变化,形态参数的样点间差异也因季节的不同而异;轮虫的各形态参数均随pH的升高而延长,随过滤水体中叶绿素a含量和水体中总叶绿素a含量的升高而缩短;体长和体宽均随晶囊轮虫密度的升高而增大。汀棠湖中,短后棘刺型个体出现于6月至10月,而长后棘刺型个体仅出现于6月至8月;2种形态型个体同时存在时,长后棘刺型个体在种群中所占比例始终高于短后棘刺型个体,并与水温、水体中总叶绿素a含量以及枝角类、桡足类和晶囊轮虫的密度均呈显著的正相关;2样点轮虫短后棘刺型个体各形态参数均随季节显著变化;长后棘刺型个体在1号样点内仅前外棘刺和后棘刺长有显著变化,而2号样点内仅体长、体宽和后棘刺长有显著变化;2样点内长后棘刺型个体的体长、体宽以及前外棘刺和后棘刺均显著长于短后棘刺型个体;同一形态型个体形态参数的样点间差异因季节的不同而异;短后棘刺型个体的前外棘刺和后棘刺长均与水体中总叶绿素a含量呈显著的正相关,后者与过滤水体中叶绿素a含量和枝角类密度也呈显著的正相关;长后棘刺型个体的后棘刺长与桡足类密度呈显著正相关,前内棘刺长与水温、溶氧量和晶囊轮虫密度呈显著负相关。两湖泊内剪形臂尾轮虫的后棘刺长和体长之间均呈显著的正相关,提示其后棘刺的延长并不是对体长的异速增长。     

基于rDNA ITS序列探讨粉煤灰污染对萼花臂尾轮虫种群遗传多样性的影响

本文通过rDNA ITS序列测定和分析, 比较研究了粉煤灰污染水体(灰湖)与未污染水体(凤鸣湖和汀棠湖)萼花臂尾轮虫(Brachionus calyciflorus)的种群遗传多样性。结果表明, 灰湖萼花臂尾轮虫仅有1个姐妹种, 而两对照湖泊均存在2个姐妹种, 粉煤灰污染使萼花臂尾轮虫种复合体内姐妹种数量减少。就三个湖泊共有的姐妹种I而言, 灰湖轮虫种群单倍型多样性最低(h= 0.9516), 核苷酸多样性(π = 0.0066)低于汀棠湖(π = 0.0073)却高于凤鸣湖种群(π = 0.0052), 粉煤灰污染对萼花臂尾轮虫种群遗传多样性表现出一定的破坏作用; 凤鸣湖轮虫种群核苷酸多样性最低, 可能是由于水体中晶囊轮虫(Asplanchna)和桡足类动物捕食压力较高以及周围农田污染的联合作用造成的。三湖泊间, 仅汀棠湖与凤鸣湖轮虫种群间基因交流值较低(N_m = 1.95), 种群分化指数较高(F_st = 0.11358); 而灰湖与汀棠湖以及灰湖与凤鸣湖轮虫种群间基因交流水平较高(N_m均大于4), 种群分化指数较低(F_st分别为0.03535和0.00276)。AMOVA分析显示汀棠湖与凤鸣湖轮虫种群间变异所占比例较高, 为12.87%; 而灰湖与汀棠湖以及灰湖与凤鸣湖轮虫种群间变异所占比例较低, 分别为3.78%和2.78%。粉煤灰污染水体与未污染水体萼花臂尾轮虫种群间保持较高水平的基因交流, 种群分化不明显。     

基于16S rDNA序列探讨十种臂尾轮虫的系统关系和分类地位

通过对角突臂尾轮虫、尾突臂尾轮虫、裂足臂尾轮虫、剪形臂尾轮虫、方形臂尾轮虫、壶状臂尾轮虫、红臂尾轮虫、镰肜臂尾轮虫、萼花臂尾轮虫和十指臂尾轮虫等十种臂尾轮虫和大肚须足轮虫的线粒体16s rDNA进行扩增和序列测序,结合Genebank 中十指臂尾轮虫的16S rDNA序列,使用MAGE软件构建了NJ(neighbor-joining method)树,使用mrbayes软件构建了贝叶斯树,探讨了十种臂尾轮虫之间的系统关系.结果表明,本研究所涉及的轮虫16S rDNA序列差异百分比均值为14.6%,可作为分子标记应用于轮虫属内种间系统关系研究;系统树均支持将十指臂尾轮虫、裂足臂尾轮虫隶属于臂尾轮属;壶状臂尾轮虫和红臂尾轮虫是两个独立的种.此外,还依据16S rDNA序列变异百分比推测了十种臂尾轮虫的分化时间.     

rDNA ITS sequences among morphotypes of Keratell cochlearis, Keratell quadrata and Brachionus forficula (Rotifera)

Morphological variation in rotifers is affected by environmental conditions, making it hard to identify some rotifer taxa. We examined the rDNA ITS sequences of 10 unspined (KCU1-KCU10) and 17 spined (KCS1-KCS17) Keratell cochlearis clones, 26 two-spined (KQT1-KQT26), 18 single-spined (KQS1-KQS18) and 9 unspined (KQU1-KQU9) K. quadrata clones, and 17 long-spined (BL1-BL17) and 11 short-spined (BS1-BS11) Brachionus forficula clones collected from Lake Tingtang in Wuhu city, China. Molecular phylogenetic trees were constructed by neighbor-joining, maximum-likelihood, maximum parsimony, and Bayesian inference methods using B. calyciflorus as an outgroup. The K. cochlearis clones included 20 haplotypes, the K. quadrata clones included 37 haplotypes, and the B. forficula clones included 25 haplotypes. Different morphotypes of each rotifer species had shared haplotypes. Sequence divergences were 0.1-8.9% among different K. cochlearis haplotypes, and 8.1-8.9% between KCHAP1 (KCU1 and KCU10), KCU3, KCU4 and KCU6, and the other haplotypes. Sequence divergences were 0.1-14.5% among different K. quadrata haplotypes, and 11.9-14.5% between KQS17 and the other haplotypes. Sequence divergences were 0.1-11.7% among different B. forficula haplotypes, 11.0-11.7% between BL15 and the other haplotypes, 9.3-10.1% between BS3 and the other haplotypes, and 11.7% between BL15 and BS3. The four phylogenetic trees all supported that KCHAP1, KCU3, KCU4, KCU6 and the other 16 haplotypes among the 20 K. cochlearis haplotypes, KQS17 and the other 36 haplotypes among the 37 K. quadrata haplotypes, and BL15, BS3 and the other 23 haplotypes among the 25 B. forficula haplotypes all belonged to their own isolated clades. The morphological variation of the three rotifer species was attributed mainly to phenotypic plasticity.     

Spine development in the rotifer Keratella cochlearis: induction by cyclopoid copepods and Asplanchna

SUMMARY. 1. Spine development in a clone of unspined Keratella cochlearis was promoted by a water soluble factor released by the copepods Tropocyclops prasinus and Mesocyclops edax and by the predatory rotifer Asplanchna priodonta. Between 9 and 55% of K. cochlearis populations cultured in predator-conditioned media responded to the inducing chemical.
2. The K. cochlearis form possessing a posterior spine of medium length was much less susceptible to predation by small Tropocylops and Asplanchna than the form lacking the posterior spine. These predators consistently selected the unspined form over the spined form when offered equal densities of each. However, both spined and unspined forms were equally susceptible to predation by large Mesocyclops.
3. The phenotypic variation of offspring produced from induced unspined Keratella females encompassed much of the variation reported for the taxon in North America. These morphotypes are similar to the variation of forms reported for the K. cochlearis tecta series known from Europe. We suggest that the North American morphotypes be identified in studies of this species because the presence of the posterior spine can greatly affect predator selectivity.     

Comparisons of phylogenetic hypotheses among different data sets in dwarf dandelions (Krigia,Asteraceae): Additional information from internal transcribed spacer sequences of nuclear ribosomal DNA

The internal transcribed spacer (ITS) region of the 18 S–25 S nuclear ribosomal DNA repeat was sequenced from 19 populations of the tribeLactuceae, including all species of dwarf dandelion (Krigia) and five outgroup genera. The incidence of length changes and base substitutions was at least two times higher for ITS 1 than ITS 2. Interspecific sequence divergence withinKrigia averaged 9.62% (1.61%–15.19%) and 4.26% (0%–6.64%) in ITS 1 and ITS 2, respectively. Intergeneric sequence divergence ranged from 15.6% to 44.5% in ITS 1 and from 8.0% to 28.6% in ITS 2. High sequence divergence and homoplasy among genera of tribeLactuceae suggest that the phylogenetic utility of ITS sequence data is limited to interspecific studies or comparisons among closely related genera. Trees generated from ITS sequences are essentially identical to those from restriction site comparisons of the entire nuclear ribosomal (nr) DNA region. The degree of tree resolution differed depending on how gaps were treated in phylogenetic analyses. The ITS trees were congruent with the chloroplast DNA and morphological phylogenies in three major ways: 1) the sister group relationship betweenKrigia andPyrrhopappus; 2) the recognition of two monophyletic sections,Krigia andCymbia, in genusKrigia; and 3) the monophyly of theK. occidentalis-K. cespitosa clade in sect.Cymbia. However, the two nrDNA-based trees are not congruent with morphology/chloroplast DNA-based trees for the interspecific relationships in sect.Krigia. An average of 22.5% incongruence was observed among fourKrigia data sets. The relatively high degree of incongruence among data sets is due primarily to conflict between trees based on nrDNA and morphological/cpDNA data. The incongruence is probably due to the concerted evolution of nrDNA repeating units. The results fromKrigia and theLactuceae suggest that nrDNA data may have limited utility in phylogenetic studies of plants, especially in groups which exhibit high levels of sequence divergence. Our combined phylogenetic analysis as a total evidence shows the least conflict to each of the individual data sets.     

九莲塘萼花臂尾轮虫种复合体结构的快速变化及其生态学机制

自2011年7月16日起,通过每周1次的轮虫采集、实验室克隆培养、DNA提取、COI基因扩增、序列测定和分析,研究了九莲塘水体中萼花臂尾轮虫种复合体结构的快速变化,并在28 ℃和32 ℃及1.0×10⁶、3.0×10⁶和5.0×10⁶ cells·mL^-1斜生栅藻密度下研究了轮虫两姊妹种的适合度特征.结果表明: 所得4批35条COI基因部分序列共定义22个单倍型,其中共享单倍型3个.基于COI基因部分序列构建的系统发生树将22个单倍型聚合为2个支系(支系Ⅰ和支系Ⅱ);支系Ⅰ和支系Ⅱ间的序列差异百分比为14.8%～15.6%,它们应属不同的姊妹种(姊妹种Ⅰ和姊妹种Ⅱ).姊妹种Ⅱ的相对丰度较低(仅占1/35),出现时间较短(仅在第2批次出现);姊妹种Ⅰ种群内,3个共享单倍型所代表的克隆均存在重叠现象,而其他克隆存在替代现象.三因素方差分析表明,温度对轮虫的净生殖率、种群内禀增长率和后代混交率有显著影响,食物密度对轮虫的平均寿命、净生殖率和种群内禀增长率有显著影响,姊妹种对轮虫的平均寿命、种群内禀增长率和后代混交率有显著影响,温度和姊妹种的交互作用对轮虫的净生殖率和种群内禀增长率有显著影响,温度和食物密度的交互作用对轮虫的后代混交率有极显著影响,食物密度和姊妹种的交互作用对轮虫种群内禀增长率有显著影响.姊妹种Ⅰ的种群内禀增长率显著高于姊妹种Ⅱ,平均寿命和后代混交率均显著短于或低于姊妹种Ⅱ.     

Polymorphism in stem females and successive parthenogenetic generations in Brachionus calyciflorus Pallas

We have described the polymorphism in the hatchlings of resting eggs and the morphological variations between the stem females hatched from resting eggs and their successive parthenogenetic generations in the rotifer Brachionus calyciflorus. We hatched resting eggs of B. calyciflorus in two different culture mediums: unconditioned medium (IOM) and Asplanchna-conditioned (ACM). The hatching rate of resting eggs in IOM and ACM were 32.5% and 28.5%, respectively, and showed no significant difference. Stem females hatching from these resting eggs had three morphotypes (unspined, single short-spined, and two short-spined) and over 80% of these females were spineless ones. Moreover, the frequency of each morphotype stem females showed similar tendency in IOM and ACM. The production of a variety of morphotypes among stem mothers—rather than all unspined as has been previously reported for this species—may be regarded as a form of bet-hedging in this population of B. calyciflorus. Phenotypic changes in morphology between the stem females hatched from resting eggs and their successive parthenogenetic generations in B. calyciflorus were found. The possible mechanisms, responsible for high proportion of spineless phenotype at early generations from resting eggs and increased spined phenotype in successive parthenogenetic generations, were discussed in the article.     

枇杷属内栽培种和部分野生种ITS序列分析

对不同栽培区的25种普通枇杷品种以及7种枇杷属野生种的ITS序列进行扩增并测序,采用邻接法和最大简约法进行系统发育树的构建并对枇杷属内不同种间的遗传关系进行了分析。结果表明:枇杷属植物ITS序列ITS1+5.8S rDNA+ITS2总长度为592 bp或594 bp,长度变化发生在ITS2。所有样本的ITS1和5.8S rDNA长度一样,都是223 bp和168 bp;而ITS2为201 bp或203 bp。5种枇杷属野生种的ITS序列长度为594 bp,包括栎叶枇杷、大渡河枇杷、南亚枇杷、南亚枇杷窄叶变种和大瑶山枇杷;其余2种枇杷属野生种(麻栗坡枇杷、小叶枇杷)和普通枇杷栽培种的ITS序列长度都为592 bp。所有样本ITS序列的GC含量为64.2%~64.5%,其中ITS1为64.1%~65.5%,ITS2为68.1%~72.6%。对所有样本的ITS序列比对产生44个可变位点,其中38个为简约信息位点,其中11个位于ITS1,5个位于5.8S rDNA,22个位于ITS2。最大的种间序列差异为7.7%,最小的种间差异发生在麻栗坡枇杷和小叶枇杷之间,仅为0.2%。普通枇杷种内的ITS序列差异很低,25种普通枇杷栽培种之间的序列差异为0~1.5%。所研究的枇杷属植物可分为3个分支。分支Ⅰ包括所有普通枇杷品种,分支Ⅱ包含5种野生枇杷种,包括栎叶枇杷、大渡河枇杷、南亚枇杷、南亚枇杷窄叶变种和大瑶山枇杷;分支Ⅲ由2个野生枇杷种(麻栗坡枇杷、小叶枇杷)组成。该研究结果表明ITS序列对枇杷种间鉴定和系统发育分析具有一定意义,但对普通枇杷栽培种间的鉴定作用不大。     

GENETIC VARIATION AND POLYMORPHISM IN THE INDUCIBLE SPINES OF A MARINE BRYOZOAN

Of particular value in understanding the evolution of genotypes with broad phenotypic ranges (phenotypic plasticity) are the few examples of organisms with adaptive plasticity, such as those that develop a defensive morphology in response to cues from predators. We know little about the heritability of inducible defensive characters or the range of phenotypes available for selection to act on in the field. Membranipora membranacea is a colonial marine bryozoan that produces spines within two days of exposure to waterborne predator extracts. Surveys done in 1993 and 1995 showed that the population at Friday Harbor Labs, Washington, was polymorphic for inducible spine type and was composed of a constitutively spined type that produced spines in the absence of a predator stimulus, an unspined phenotype that did not produce spines irrespective of a stimulus, and an inducibly spined phenotype that would produce spines if exposed to the appropriate cue. In 1995, the frequencies of these types were determined through a laboratory common-garden experiment; 178 colonies were cultured from metamorphosis through approximately 30 days and then exposed to the cue. The inducible type was the dominant, comprising 80.3% of the population. The constitutive type made up 6.2% of the population, and the remaining 13.4 % was the unspined type. The frequency of the three types was similar to a preliminary trial of the experiment run in 1993. Experiments also showed that the lengths of the spines of the inducible type varied continuously among genotypes. To assess causes of variation in the inducible spine response and its clonal heritability, 16 clones were subdivided and grown in a common environment and exposed to a single dosage of spine inducing substance (SIS). Spine length showed high clonal heritability. The range of colony responses from a single environment varied from relatively unresponsive to highly responsive colonies with a very low threshold of response. Norms of reaction were quantified for spine lengths of inducible genotypes originating from two field environments by testing them in a concentration series of SIS. Both spine length and spine type varied with concentration of inducer. Within a clone, colonies were more likely to produce membranous spines than corner spines at higher concentrations. At low concentrations, only straight spines were produced. This study showed that populations of M. membranacea at Friday Harbor are a mix of inducible, nonspined and constitutively spined individuals. Even the inducible individuals showed high heritable variation in the length of spine activated, suggesting that there is considerable scope for the evolution of this character. A norm-of-reaction experiment further showed that the type of spine produced, membranous or corner, varied with the concentration of the cue. Factors maintaining the polymorphism and the broad range of genotypes could include high costs of defending the spined types coupled with a shifting biotic regime.     

Effect of Temperature on Fecundity,Life Span and Morphology of Long‐ and Short‐Spined Clones of Brachionus caudatus f. apsteini (Rotifera)

We investigated the effect of temperature (20, 25 and 30 °C) on fecundity, life span and morphology of the rotifer Brachionus caudatus f. apsteini. For each temperature, short posterior‐spined and long posterior‐spined clones of B. caudatus f. apsteini were individually cultured for up to six generations. The rotifers were fed Chlorella sp. at a density of 1 × 10⁶ cells ml^–1. Morphometric data (body size and spine length) were collected. Total number of offspring producing by a single female per life cycle at high temperature was higher than at low temperature. The duration of juvenile period, reproductive period, post‐reproductive period and life span of both clones of B. caudatus f. apsteini decreased with increasing temperature. All offspring of short posterior‐spined clone produce posterior spines at 20 and 25 °C, with an average length of 19.8 ± 6.6 and 11.9 ± 2.6 μm, respectively. In contrast, they cannot develop posterior spines at 30 °C, at which the average length of the posterior spine remnant was 6.4 ± 1.3 μm. On the other hand, all offspring of long posterior‐spined clone have long posterior spines with average lengths of 36.8 ± 6.1, 36.3 ± 5.2 and 36.6 ± 6.2 μm at 20, 25 and 30 °C, respectively. This study indicated that the production of posterior spines can be induced by low temperature and that short posterior‐spined and long posterior‐spined clones are genetically different. (© 2008 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)