Halbach Udo
Zoologisches Institut der Universität München, München, Deutschland.
Oecologia. 1971 Sep;6(3):267-288. doi: 10.1007/BF00344919.
In the rotifer Brachionus calyciflorus there occurs a cyclomorphic variation concerning the absence or the presence of a pair of postero-lateral spines. If present, these spines may vary up to lorica length. The production of these spines is induced by starvation, low temperature, or a peptide substance released into the medium by the predacious rotifer Asplanchna. The first two factors apparently induce spines only up to a relative short length, while extremely long spines appear to be induced only by the Asplanchna substance. 1. Existing hypotheses with regard to possible adaptive values of the cyclomorphic spine production are discussed and rejected with the exception of two, which are tested by laboratory experiments. 2. The hypothesis of Erman, that spined Brachionus calyciflorus swim more slowly and filter the nourishment more effectively as a consequence of the resistance of the spines to friction, is disproved experimentally (see Table 1 and 2). 3. The hypothesis of Beauchamp and of Gilbert, that spines form a protection against the predacious Asplanchna, is tested in detail. a) The components of the predator-prey relationship of Asplanchna and the influences of the spines on these components are studied in short term experiments (stopped after the first catch; see Fig. 2 and Table 3): The number of accidental collisions with Asplanchna is not influenced by spine length. However, by increasing spine length the rate of catches per collision and the rate of swallowed prey per catch decrease, and the time spent in swallowing the prey increases. The timespan from the start of the experiment to the moment when the first prey is just being swallowed shows a marked increase with growing spine length (see Table 3: g). b) In experiments of longer periods of exposition (up to one hour; see Table 5) the long-spined B. calyciflorus are again nearly completely unassailable. In these cases the advantage of short-spined animals (compared with spineless ones) is reduced. The reason for this is a more rapid saturation of the Asplanchna by unarmed prey. This causes a quicker relaxation of the predator's activities (decrease of the swimming speed, number of prey caught per collision, and number of prey swallowed per catch). In Asplanchna feeding on short-spined prey, saturation is slower but at the same time the predator's activities decrease more slowly, too (see Fig. 4). By extending the periods of exposition the survival chances of spined and spineless animals become more similar. c) In the presence of spineless animals (Brachionus rubens) as an alternative prey, the protection of the spines (in relation to spineless controls) becomes more important, because in this case the unarmed alternative prey makes a higher proportion of the predator's diet. This is true over a wide range of predation pressure (see Tables 6 and 7). d) Protective spinesara also present in juvenile B. calyciflorus. However, the juveniles are in all of their categories of spine length more sensitive to predator attacks than are corresponding adult females. The reason for this is the smaller overall body size of these animal (Tables 3, 4, 5, 6, Fig. 4). 4. The connexions between the inductive factors and the adaptive values of the cyclomorphic spine production are reviewed and discussed.
在萼花臂尾轮虫中,存在一种关于一对后侧棘有无的轮虫形态变异。如果有这些棘,其长度可能会变化至与兜甲长度相当。这些棘的产生是由饥饿、低温或捕食性轮虫晶囊轮虫释放到培养基中的一种肽物质诱导的。前两个因素显然仅能诱导出相对较短长度的棘,而极长的棘似乎仅由晶囊轮虫物质诱导产生。1. 除了两个通过实验室实验进行检验的假设外,关于轮虫形态棘产生可能的适应性价值的现有假设均被讨论并否定。2. 埃尔曼的假设,即有棘的萼花臂尾轮虫由于棘对摩擦的阻力而游动更慢且更有效地过滤营养物质,已通过实验被证伪(见表1和表2)。3. 博尚和吉尔伯特的假设(即棘形成了对捕食性晶囊轮虫的保护)得到了详细检验。a) 在短期实验(首次捕获后停止;见图2和表3)中研究了晶囊轮虫捕食关系的组成部分以及棘对这些组成部分的影响:与晶囊轮虫的意外碰撞次数不受棘长度的影响。然而,随着棘长度的增加,每次碰撞的捕获率以及每次捕获所吞咽猎物的速率降低,并且吞咽猎物所花费的时间增加。从实验开始到首次吞咽猎物的时间跨度随着棘长度的增加而显著增加(见表3:g)。b) 在较长暴露时间(长达一小时;见表5)的实验中,长棘的萼花臂尾轮虫再次几乎完全无法被攻击。在这些情况下,短棘动物(与无棘动物相比)的优势降低。原因是无武装的猎物使晶囊轮虫更快地达到饱和。这导致捕食者活动更快地放松(游泳速度、每次碰撞捕获的猎物数量以及每次捕获吞咽的猎物数量减少)。以短棘猎物为食的晶囊轮虫,达到饱和的速度较慢,但同时捕食者活动的降低也较慢(见图4)。通过延长暴露时间,有棘和无棘动物的生存机会变得更加相似。c) 在有无棘动物(红臂尾轮虫)作为替代猎物的情况下,棘的保护作用(相对于无棘对照)变得更加重要,因为在这种情况下,无武装的替代猎物在捕食者的食物中占更高比例。在广泛的捕食压力范围内都是如此(见表6和表7)。d) 有保护作用的棘也存在于幼年萼花臂尾轮虫中。然而,这些幼体在所有棘长度类别中都比相应的成年雌性对捕食者攻击更敏感。原因是这些动物的整体体型较小(表3、4、5、6,图4)。4. 回顾并讨论了诱导因素与轮虫形态棘产生的适应性价值之间的联系。