Effect of different densities of live and dead Chlorella vulgaris on the population growth of rotifers Brachionus calyciflorus and Brachionus patulus (Rotifera)

In order to maintain rotifer populations during periods of low algal production, it is necessary to offer alternate diets, some of which include forms of preserved algae. The present work is based on the effect of live and dead Chlorella vulgaris on the population growth of Brachionus calyciflorus and Brachionus patulus. The experimental design consisted of three algal levels (0.5 x 10(6), 1.5 x 10(6) and 4.5 x 10(6) cells ml-1) offered in three forms (living, frozen and heat-killed). The maximal population density values for B. calyciflorus ranged from 55 +/- 1 ind. ml-1 (at 0.5 x 10(6) cells ml-1) to 471 +/- 72 ind. ml-1 (at 4.5 x 10(6) cells ml-1) with live Chlorella, but was much lower (6 +/- 1 to 26 +/- 6 ind. ml-1) with frozen or heat-killed alga under comparable food levels. However, the maximum population density of B. patulus under live or or heat-killed Chlorella was similar at comparable algal levels but when offered frozen algae it was four times less. The highest mean peak population density was 1,277 +/- 83 ind. ml-1 under 4.5 x 10(6) cells ml-1. The rate of population increase for B. calyciflorus varied from 0.50 to 0.79 using live Chlorella, but under comparable conditions, this range was lower (0.21 to 0.31) for B. patulus. Results have been discussed in light of possible application for aquaculture.     

Combined effect of concentrations of algal food (Chlorella vulgaris) and salt (sodium chloride) on the population growth of Brachionus calyciflorus and Brachionus patulus (Rotifera)

Salinity is an important variable influencing the density and diversity of rotifers. Studies on salt tolerance of rotifers have so far concentrated on euryhaline species while very little information is available on non-euryhaline taxa. In the present work, we have evaluated the combined effects of Chlorella vulgaris and sodium chloride on the population growth of two freshwater rotifers B. calyciflorus and B. patulus. A 24 hr acute tolerance test using NaCl revealed that B. calyciflorus was more resistant (LC50 = 3.75 +/- 0.04 g l-1) than B. patulus (2.14 +/- 0.09 g l-1). The maximal population density (mean +/- standard error) for B. calyciflorus in the control at 4.5 x 10(6) cells ml-1 (algal level) was 80 +/- 5 ind. ml-1, which was nearly a fifth of the one for B. patulus (397 +/- 7 ind. ml-1) under comparable conditions. Data on population growth revealed that regardless of salt concentration, the density of B. calyciflorus increased with increasing food levels, while for B. patulus, this trend was evident only in the controls. Regardless of salt concentration and algal food level, the day of maximal population density was lower (4 +/- 0.5 days) for B. calyciflorus than for B. patulus (11 +/- 1 day). The highest rates of population increase (r values) for B. calyciflorus and B. patulus were 0.429 +/- 0.012 and 0.367 +/- 0.004, respectively, recorded at 4.5 x 10(6) cells ml-1 of Chlorella in the controls. The protective role of algae in reducing the effect of salt stress was more evident in B. calyciflorus than B. patulus.     

温度和食物浓度对三品系萼花臂尾轮虫实验种群动态的影响

应用个体培养方法,研究了温度(20、25和30℃)和藻类食物浓度(1．5×10^6、3．0×10^6、6．0×10^6和9．0×10^6cells·ml^-1)对青岛、芜湖、广州三品系萼花臂尾轮虫种群动态的影响．结果表明,温度仅对轮虫的世代时间和种群内禀增长率有显著影响,而品系对所有生命表参数均无显著影响．轮虫种群的内禀增长率随培养温度的升高而增大,世代时间则随培养温度的升高而缩短．食物浓度仅对轮虫的生命期望值和平均寿命有显著影响,品系对轮虫的净生殖率、世代时间、生命期望值和平均寿命也有显著影响．三品系间,以广州品系轮虫的净生殖率、世代时间、生命期望值和平均寿命最大,芜湖品系最短．当食物浓度为3．0×10^6cells·ml^-1时,轮虫的生命期望值和平均寿命最长,9．0×10^6cells·ml^-1时最短．各品系轮虫的净生殖率、世代时间、总生殖率、生命期望值和平均寿命均随培养温度的升高而减小,广州品系的净生殖率除外．轮虫种群的内禀增长率和广州品系轮虫的总生殖率则随培养温度的升高而增大．青岛和广州品系轮虫的各生命表参数,均与食物浓度呈曲线相关,但芜湖品系仅世代时间、平均寿命和生命期望值随食物浓度的增大而缩短．     

Life history characteristics of three types of females in Brachionus calyciflorus Pallas (Rotifera) fed different algae

This study describes the life history characteristics of amictic, unfertilized mictic and fertilized mictic females of the rotifer Brachionus calyciflorus cultured individually on two different algae at 0.1 mg ml^–1 food concentration and 27 °C. The duration of the juvenile period of amictic females was significantly shorter on Chlorella pyrenoidosa Chick than on Scenedesmus obliquus Kütz or both algae together. The duration of the juvenile period of unfertilized mictic females was significantly longer, and the number of eggs produced by amictic females was significantly larger on Chlorella pyrenoidosa than on S. obliquus. When fed the same type of alga, the duration of the juvenile period of the fertilized mictic females was the longest among the three types of females, and the durations of the reproductive period of the amictic females and the post-reproductive period of the fertilized mictic females were longer than, or equal to those of the other two types of females, respectively. The number of eggs produced by an unfertilized mictic female was the largest among the three types of females, and that of amictic females was larger than or equal to that of fertilized mictic females, depending on the type of diet.     

三氯杀螨醇浓度和食物密度对萼花臂尾轮虫种群增长的影响

采用群体累积培养方法,研究了不同浓度(0.03、0.3、3.30和300 μg·L-1)的三氯杀螨醇和不同食物密度(3.0和5.0×106cells·ml-1)的斜生栅藻(Scenedesmus obliquus)对萼花臂尾轮虫(Brachionus calyciflorus)种群增长影响.结果表明:藻类食物密度、三氯杀螨醇浓度以及二者的交互作用对轮虫种群增长率均有显著影响(P＜0.05);藻类食物密度和三氯杀螨醇浓度对轮虫最大种群密度也有显著影响(P＜0.01),但二者的交互作用对其却无显著作用(P＞0.05);与空白对照组相比,在3.0×106cells·ml-1藻密度下.0.03-30μg·L-1的三氯杀螨醇显著提高了轮虫种群增长率,3μg·L-1的三氯杀螨醇显著降低了轮虫的最大种群密度,而300μg·L-1的三氯杀螨醇则极显著提高了轮虫的最大种群密度;在5.0×106cells·ml-1藻密度下,三氯杀螨醇对萼花臂尾轮虫种群增长率和最大种群密度均无显著影响;高密度的藻类食物降低了0.03-30μg·L-1和300μg·L-1的三氯杀螨醇分别对萼花臂尾轮虫种群增长率和最大种群密度所具有的促进作用,以及3μg·L-1的三氯杀螨醇对轮虫最大种群密度所具有的抑制作用.     

Comparative population growth and life table demography of the rotifer Asplanchna girodi at different prey (Brachionus calyciflorus and Brachionus havanaensis) (Rotifera) densities

Population growth and life table demography of the predatory rotifer A. girodi using spineless Brachionus calyciflorus and spined Brachionus havanaensis as prey at densities of 1, 2, 4 and 8 ind. ml^–1 at 25°C were studied. Regardless of the prey species, the population of A. girodi increased with increasing availability of Brachionus in the medium. At any given prey density, A. girodi fed B. calyciflorus showed consistently better growth than when fed B. havanaensis. The maximum population densities of A. girodi varied from 0.28 to 1.8 ind. ml^–1 depending on the prey species and the density. The rate of population increase observed in population growth studies varied from 0.17 to 0.43 day^–1 when fed B. calyciflorus and 0.09 to 0.27 day^–1 when fed B. havanaensis. Male population of A. girodi was closely related to female density. The lowest average lifespan was observed for A. girodi when fed B. havanaensis at 1 ind. ml^–1, while the converse was the case when fed B. calyciflorus at comparable prey concentration. Net reproductive rates varied from 16 to 26 offspring female^–1 lifespan^–1 depending on the prey species and concentration. Generation time of A. girodi decreased with increasing food concentrations for both the prey species. The rates of population increase obtained from life table demography were lower for A. girodi when fed B. havanaensis than when fed B. calyciflorus.     

Nomenclatural consequences of some recent studies on Brachionus plicatilis (Rotifera,Brachionidae)

Recent studies on morphology, karyotype, genetics including allozyme constitution and reproductive behaviour of S- and L-type B. plicatilis showed that these types are best treated as different species. A reexamination of existing available names revealed B. plicatilis O. F. Müller, 1786 and B. rotundiformis Tschugunoff, 1921 as the correct names for the L- and S-type, respectively.     

Effects of three organophosphorus pesticides on population growth and sexual reproduction of rotifer Brachionus calyciflorus Pallas

Effects of organophosphorus pesticides including dichlorvos, triazophos and chlorpyrifos on population growth and sexual reproduction of freshwater rotifer Brachionus calyciflorus were studied by 3-d population growth and 4-d resting egg production tests. The results showed that all the three organophosphorus pesticides influenced significantly the population growth rate, the ratio ovigerous females/non-ovigerous females in the rotifer populations and the resting egg production of the rotifers. Both dichlorvos and chlorpyrifos influenced markedly the mictic rate of the rotifers, but triazophos did not. Compared to the controls, both dichlorvos at 10.0–1000.0 μg/L and chlorpyrifos at 0.01–100.0 μg/L increased the population growth rate, but the reverse was also true for dichlorvos and triazophos both at 10000.0 μg/L. Chlorpyrifos at 10000.0 μg/L made the rotifers dead after 24-hr exposure. Dichlorvos and triazophos both at 10000.0 μg/L, and chlorpyrifos at 1000.0 μg/L all increased the ratio ovigerous females/non-ovigerous females. Both dichlorvos at 10000.0 μg/L and chlorpyrifos at 0.1–100.0 μg/L increased the mictic rate. Dichlorvos at 10.0 μg/L and 100.0 μg/L, and triazophos and chlorpyrifos both at 0.1–100.0 μg/L increased the resting egg production. Both population growth rate and ratio ovigerous females/non-ovigerous females are suitable endpoints for assessing the effects of dichlorvos, triazophos and chlorpyrifos, and mictic rate is a suitable endpoint for monitoring the effects of dichlorvos and chlorpyrifos on the reproduction of the rotifers. Both population growth rate and ratio ovigerous females/non-ovigerous females are more sensitive to dichlorvos and chlorpyrifos than mictic rate.     

Demographic parameters of Brachionus patulus Muller (Rotifera) exposed to sublethal DDT concentrations at low and high food levels

The combined effects of sublethal levels of DDT (0, 15, 30, 45, and 60 µg·l^–1) and food (Chlorella at 1 and 3 × 10⁶ cells ml^–1) on the demography of the rotifer Brachionus patulus were studied. The average lifespan, life expectancy, net reproductive rate (R_o) and intrinsic rate of increase (r) were significantly greater at the higher food level, but declined with increasing DDT concentration. A significant DDT food interaction was manifested in the toxicity-mitigating action of the higher food density at higher DDT levels. The r values were negative at DDT levels 30 µg · l^–1 and above, only when food density was low. The effective DDT concentration at which a given parameter value was reduced to 50% of that in the controls (EC50) was lower at lower food levels for both survivorship and reproduction, and lower for reproduction than for survivorship at either food level.     

镜湖萼花臂尾轮虫夏季种群等位酶分析以及克隆间的生殖差异

应用聚丙烯酰胺凝胶垂直板电泳技术和特异性染色方法,对采自芜湖市镜湖的萼花臂尾轮虫(Brachionuscalyciflorus)夏季种群内的16个克隆进行了等位酶分析。在检测的6种等位酶系统中,依据葡萄糖磷酸异构酶(GPI)和苹果酸脱氢酶(MDH)的酶谱在克隆间所存在的明显差异,将镜湖萼花臂尾轮虫夏季种群内的16个克隆划分为4个互不相同的克隆群。对由4个克隆群中分别随机选取的1个克隆(共4个克隆,分别命名为克隆A、克隆B、克隆C和克隆D)在4种斜生栅藻(Scenedesmusobliquus)密度(1.0,2.0,4.0和8.0×106cells/ml)下的孤雌生殖和有性生殖所作的研究发现,食物密度、克隆以及两者间的交互作用对轮虫的种群瞬时增长率、种群中的混交雌体百分率、混交雌体受精率和休眠卵产量等4项指标均有显著影响。4克隆中,克隆D的这4项指标均最高,而克隆B的种群瞬时增长率以及克隆A的其他3项指标均最低。克隆B的种群瞬时增长率、种群中的混交雌体百分率和混交雌体受精率均与食物密度间无显著的相关性;克隆A和C的种群瞬时增长率与食物密度间呈曲线相关,而其种群中的混交雌体百分率和混交雌体受精率均与食物密度间无显著的相关性;克隆D的种群瞬时增长率、种群中的混交雌体百分率和混交雌体受精率均与食物密度间呈曲线相关。这些结果表明:镜湖萼花臂尾轮虫夏季种群内存在“克隆共存”现象,且有些克隆间在生化遗传和生态特征方面具有明显的差异     

镜湖萼花臂尾轮虫种群内不同基因型克隆的生活史特征

应用生命表统计学等方法,比较了15、20、25和30℃4个温度下镜湖萼花臂尾轮虫种群内13个不同基因型克隆的生命表统计学参数和后代混交雌体百分率等生活史特征．结果表明：轮虫的世代时间、平均寿命、出生时的生命期望值、内禀增长率、净生殖率和后代混交雌体百分率对升高温度的反应均因克隆的不同而存在差异;温度、克隆及两者间的交互作用对其也都具有极显著影响．表明水环境在温度等方面所具有的时间异质性可能是镜湖萼花臂尾轮虫种群丰富的遗传变异得以维持的原因之一;自然选择出现在共存的克隆中,但其强度较低或作用时间较短是克隆共存的重要原因;遗传漂变可能在种群的基因型频率变化中起重要作用．不同基因型轮虫克隆的存在对其种群在水环境中的持续存在具有重要意义．     

镜湖萼花臂尾轮虫夏季种群内不同克隆的生活史特征

应用生命表统计学等方法对镜湖萼花臂尾轮虫夏季种群内4个在生化遗传特征上互不相同的克隆（克隆A、B、c和D）在4种温度（15℃、20℃、25℃和30℃）下的生活史特征及其对升高的温度的反应进行了比较研究。结果表明,温度对萼花臂尾轮虫存活率和繁殖率的影响在不同克隆间存在着差异。20℃和25℃下,4个克隆轮虫的世代时间和净生殖率均分别无显著的差异;20℃、25℃和30℃下,4个克隆轮虫的平均寿命和出生时的生命期望亦然。其余各温度下,4个克隆间轮虫的其他生命表参数均有显著的差异。20℃、25℃和30℃下,4个克隆轮虫所产后代中的混交雌体百分率间均具有显著的差异,且克隆c轮虫所产后代中的混交雌体百分率最高。15℃下,克隆D轮虫的个体适合度最高;25℃下,克隆A和B轮虫的个体适合度较高。4个克隆轮虫的生活史特征对升高的温度的反应也存在着差异。忽略温度的影响时,4克隆间,克隆D轮虫的世代时间、平均寿命和出生时的生命期望最短,净生殖率、种群内禀增长率和个体适合度均最低;克隆C轮虫所产后代中的混交雌体百分率最高。30℃下,4个克隆轮虫的种群内禀增长率存在着差异可能是不同基因型的轮虫克隆群在种群内所占的比例不同的重要原因;而它们的个体适合度相似则可能是不同基因型的轮虫克隆群在夏季镜湖中共存的主要原因之一。     

Combined effects of food level and inoculation density on competition between Brachionus patulus (Rotifera) and the cladocerans Ceriodaphnia dubia and Moina macrocopa

Competition among cladocerans and rotifers is of considerable interest not only due to their close similarity in life history strategies, but also due to the considerable overlap they exhibit in their feeding habits. In tropical waterbodies, several genera of cladocerans, including Ceriodaphnia and Moina occur, simultaneously with rotifers. We tested over a period of 3 weeks the combined effects of food (0.5×10⁶ and 1.5×10⁶ cells ml^–1 of Chlorella) level and rotifer density on the competition between B. patulus and C. dubia and M. macrocopa using population growth experiments. For each cladoceran species we used 30 test jars of 50 ml capacity. The initial density of cladocerans was 0.2 ind ml^–1, while for B. patulus it was either 1 ind ml^–1 or 5 ind ml^–1. Neither the maximal population density nor the rate of population increase (r) of C. dubia was significantly affected by B. patulus. However, for M. macrocopa, both these variables were negatively affected by the rotifers. The combined effects of low food level and high initial density of B. patulus resulted in a 50% reduction in the peak population density of M. macrocopa. The population growth of B. patulus was negatively influenced by the presence of C. dubia and M. macrocopa. The results of the competition experiments conducted in the present study between cladocerans and rotifers suggest the existence of a more complex and delicate interaction than is generally thought.     

pH值对萼花臂尾轮虫种群增长及繁殖的影响

采用种群积累培养法,实验观察了 ｐＨ在３．５～１１．５之间（间隔 １）萼花臂尾轮虫 （Ｂｒａｃｈｉｏｎｕｓ ｃａｌｙｃｉｆｌｏｒｕｓ）种群的增长及繁殖．结果表明,该轮虫种群在ＰＨ６．５～８．５之间增 长较快, ８． ５时增长最快,即瞬时增长率 ｒ和种群密度较大和最大; ｐＨ在 ３． ５～４． ５和 ９． ５ ～１０． ５之间,种群为负增长,即 ｒ为负值; ｐＨ在 ５． ５～９． ５之间种群为正增长,即正为正 值．该轮虫存活的ｐＨ上限为１１．５,下限为３．５．在种群增长最适ｐＨ（８．５）条件下,该轮虫 的繁殖最快,即绝对带卵量最高（１３２个·ｍｌ－１）;ｐＨ在９．５时,其相对带卵量最高．为其它 ｐＨ值条件下的２～４倍．本研究结果可为淡水轮虫的大批量培养提供可靠的ｐＨ值技术指 标．     

溴氰菊酯对萼花臂尾轮虫实验种群动态的影响

采用生命表技术对暴露于不同浓度溴氰菊酯溶液中的萼花臂尾轮虫的存活和繁殖进行了研究。结果显示, 溴氰菊酯使轮虫的存活时间显著缩短, 繁殖率降低; 当溴氰菊酯浓度高达3 6 mg/L时, 轮虫的存活时间最短、繁殖率最低。与对照组相比, 除1 2 mg/L外, 其它各浓度的溴氰菊酯均使轮虫的生殖前期显著延长; 浓度为0 6和1 2 mg/L的溴氰菊酯使轮虫的生殖期显著延长, 而浓度为2 4和3 0 mg/L的溴氰菊酯却使轮虫的生殖期显著缩短; 1 2 mg/L的溴氰菊酯使轮虫的平均寿命显著延长。轮虫的生命期望、世代时间、净生殖率和种群内禀增长率均随溴氰菊酯浓度的升高而下降。当溴氰菊酯浓度升高达1 2 mg/L时, 轮虫的净生殖率开始与对照组有显著差异; 而轮虫的种群内禀增长率从溴氰菊酯浓度升高达2 4 mg/L时才开始与对照组有显著差异。在溴氰菊酯的毒性监测中, 净生殖率是比种群内禀增长率更敏感的指标。     

Isolation and purification of glutathione S-transferases from Brachionus plicatilis and B. calyciflorus (Rotifera)

1. The enzyme glutathione S-transferase (GST), a critical element in xenobiotic metabolism, was isolated from the marine rotifer Brachionus plicatilis and its freshwater congener B. calyciflorus. 2. In B. plicatilis, GST comprised 4.2% of cytosolic protein and was present as three separate isozymes with mol. wts 30,000, 31,400 and 33,700. Specific activity of crude homogenates was 56 nmol min-1 mg-1 protein, while that of affinity chromatography purified GST was 1850. 3. In B. calyciflorus, GST was present as two isozymes with mol. wts of 26,300 and 28,500, representing 1.0% of cytosolic protein. Crude GST specific activity was 1750 nmol min-1 mg-1 protein and purified was 72,400. 4. Rotifer GSTs are unusual because they are monomers whereas all other animals thus far investigated posses dimeric GSTs.     

Transgenerational and within-generational induction of defensive morphology in Brachionus calyciflorus (Rotifera): importance of maternal effect

Maternal effects on the induction of defensive morphology are regarded as an adaptive strategy. For instance, a female can transmit environmental conditions related to predation risk to offspring of the same generation or succeeding generations. However, studies have presented limited information regarding such an adaptive maternal effect on monogonont rotifers. In the present study, the maternal effects on the spine development of rotifers were evaluated using Brachionus calyciflorus-Asplanchna models. Asplanchna can release soluble kairomones into the environments, thereby inducing B. calyciflorus to form defensive morphs (e.g., elongated posterolateral spines). Our empirical data supported the hypothesis that an amictic Brachionus-experienced Asplanchna kairomones could produce offspring with longer posterolateral spine in the current or succeeding generation than those of a non-experienced amictic female. On the basis of our data, we speculated that the maternal effect of B. calyciflorus may involve the transfer and dilution of inducers (e.g., hormone) within and between generations. On one hand, our results reinforced the within-generation maternal effects on induction of defensive morphology of rotifers; on the other hand, our results enhanced transgenerational inducible defense to other planktonic taxa that have been only observed in cladocera.     

Acute toxicity and bioaccumulation of endosulfan in rotifer (Brachionus calyciflorus).

1. The acute toxicity of endosulfan was determined for the freshwater rotifer Brachionus calyciflorus. 2. The mean 24 hr LC50 value for endosulfan was 5.15 ppm with a coefficient of variation of 14.7%. 3. Rotifers were exposed at two sublethal concentrations (1.5-2.0 ppm) of endosulfan for bioaccumulation experiments, for an exposure time of 24, 48, 72 and 96 hr. The rotifers were fed with Nannochloris oculata (5 x 10(5) cell/ml). 4. The highest accumulation of endosulfan was found 24 hr after the start of the exposure to 1.5 ppm of the toxicant. A steady-state concentration in rotifer was reached between 24-48 hr, followed by a gradual decrease until 96 hr.     

邻苯二甲酸酯类物质对萼花臂尾轮虫种群增长和有性生殖的影响

以萼花臂尾轮虫(Brachionus calyciflourus)为受试动物,以17β-雌二醇(E2)为阳性对照,运用3d种群增长和4d休眠卵产量实验方法研究了不同浓度的邻苯二甲酸二丁酯(DBP)、邻苯二甲酸二异辛酯(DEHP)和邻苯二甲酸丁苄酯(BBP)对轮虫种群增长和有性生殖的影响。结果显示,与空白对照组相比,0.05-5000μg/L的E2和500μg/L的BBP显著提高了轮虫种群增长率,而0.5和5000μg/L的DBP以及0.05和0.5μg/L的BBP使轮虫种群增长率显著降低;0.5-500μg/L的E2、50和500μg/L的DBP、0.005和500μg/L的DEHP和500μg/L的BBP显著提高了轮虫的休眠卵产量,而5000μg/L的DBP和0.05μg/L的BBP却与其相反;5和500μg/L的E2、50和500μg/L的DEHP以及500μg/L的DBP和BBP均使轮虫混交率显著上升,而0.005和0.05μg/L的E2、5000μg/L的DBP和DEHP及0.5μg/L的BBP显著降低了轮虫种群中混交雌体受精率;除5000μg/L的E2显著降低轮虫种群中的混交雌体数/非混交雌体数之外,其它所有处理组均对其无显著性影响;5、500和5000μg/L的E2、5000μg/L的DBP和5-5000μg/L的DEHP均使轮虫种群中的带卵雌体数/不带卵雌体数显著升高,而BBP对其无显著影响。在实验设置浓度范围内,轮虫种群中的混交雌体数/非混交雌体数与E2浓度间、轮虫种群中的混交雌体受精率和种群增长率与DBP浓度间、轮虫休眠卵产量与DEHP浓度间、轮虫混交率与BBP浓度间均具有显著的效应-剂量关系     

种群起始密度对萼花臂尾轮虫休眠卵形成的影响

应用群体累积培养法,研究了种群起始密度对萼花臂尾轮虫休眠卵的形成及之相关的混交雌体百分率和受精率的影响,结果表明,在连续１１ｄ的培养中,轮虫休眠卵的产量和形成效率以及种群中的平均混交雌体百分率以种群起始密度为１．０－１０．０ｉｎｄ．ｍｌ＾－１组最大,２０．０ｉｎｄ．ｍｌ＾－１组次之,５０．０ｉｎｄ．ｍｌ＾－１组最小,各组间的平均混交雌体受精率则无显著差异,对轮虫休卢卵最大形成效率及其出现的时间以及