Marchenkov A V
Parazitologiia. 2001 Sep-Oct;35(5):406-28.
According to the rule of academican E. N. Pavlovskiy, any organism of host is an environment of inhabit for a parasite (Pavlovskiy, 1934). It was analysed, which "ecological niche" or microbiotop (= microhabitat) is occupied by this or that species of symbiotic (parasitic) copepods in organisms of different groups invertebrate-hosts. The assumption lying in a basis of the given analysis means that each group of hosts may give to cohabitants only certain variants of microbiotopes independently on the general morphological structure and life mode of hosts. Five types of microbiotops offered by various groups of hosts for symbiotic copepods are designated (Ta[symbol: see text] 2). 1. The body surface of benthic invertebrates as a microbiotope is characterized by conditions being little different (concerning any kind of physical and chemical influences on copepods) from those in external environment on any other substrate. Apparently a trophical dependence plays a determining role in this case. There are certain directions in a development of adaptations, which are characteristic in some extent for all water ectoparasitic crustaceans and have one functional task--to help to an ectoparasite to keep itself on a surface of host body. In the first, the maxillules and maxillipeds significantly are developed, they get a form of large claws, with which the dopepods are strongly attached on a surface of host body and have an opportunity to move on it without a danger to be washed off. In the second, the form of the body undergoes a dorso-ventral expression and expansion of prosome, forms a cephalic shield allowing to the symbiont to press itself tightly to the host body surface and to avoid the loss of host (tab. 2). In occasions, some ectoparasites stimulate the formation of galls in skin tissues of the host, that also provides the parasite with constant conditions, without any threat to lose the host. However, this phenomenon has not a wide distribution and is observed in some groups of crustacean and echinoderm hosts. 2. The narrow tubular cavities in the organism of host either they are a part of external environment (as in channel system of spongia) or a part of internal environment of organism (as channels of blood system or thin parts of a digestive system) have always rigidly limited sizes and form. Characteristics of all parasites occupying this microbiotopes are the strong transformations. They are expressed by the reduction of legs or any other appendages (frequently in a significant degree), loss of segmentation to some extent and in eruciform (or vermiform) form of a body (tab. 2). This microbiotope is occupied by an ectoparasite in one case only (Spongicola uncifer from channel system of spongia) and by endoparasites in all other cases. 3. Large cavities connected with external environment. The formations of various genesis, such as mantle cavity of molluscs, gill cavity and marsupium of crustaceans, bursal cavity of ophiuroids and branchial cavity of ascidians, concern this type of microbiotopes. All of them are characterized by the relative difference from the external environment and rather large volume (in comparison to sizes of copepods), that provides the parasites with a sufficient protection from factors of the external environment and constant source of food such as elements of host body or food's particles brought by the water flow. Morphological changes in inhabitants of the microbiotope have two directions. They practically are absent in the overwhelming majority copepods, living in the mantle of cavity of the lamellibranches. On the other hand, the inhabitants of gill cavity and marsupium of crustaceans, bursal cavities of ophiuroids and branchial cavity of ascidians are characterized by the presence of strong transformations. Usually there are expressed in a loss of segmentation to some extent, reduction of appendages and swelling of body, as in species of the genus Sphaeronella (tab. 2). Changes are also observed in the life cycle: the tendency to reduce stages of development (development of nauplii stage, which takes place under the ovarial cover). In this case the copepodid stages hatch from the ova. 4. The internal cavity of organism of host. This type of microbiotopes in different groups of the hosts is represented in a various degree. We recognise it in a coelome of polychaetes, lacunar system of molluscs, mixocoel of crustaceans, coelome of echinoderms and cavity of body in ascidians. Two basic evolutionary directions are observed in copepods occupying this microbiotope. In the first case, the parasite is not exposed to transformations and keeps the initial plan of structure as in ancestral free-living forms. In the second case the parasites are exposed to strong transformations, they either live directly in cavity's liquid, or are surrounded by a cyst (as in Cucumaricolidae). 5. Microbiotope of the last type is most specific. The simultaneous existence in two environments--external environment (environment of the second order) and internal environment (environment of the first order) leads to the complete loss of ancestral type in a structure and level of organisation. At the same time both morphological and functional division of the parasite body into two parts produces a new formation--the ectosome and endosome. In this case we deals with the phenomenon of mesoparasitism.
根据院士E. N. 巴甫洛夫斯基的理论,宿主的任何生物体都是寄生虫的栖息环境(巴甫洛夫斯基,1934年)。本文分析了不同类群无脊椎动物宿主的生物体中,各种共生(寄生)桡足类动物占据何种“生态位”或微生物群落(=微生境)。该分析的基本假设是,每类宿主给予共生者的微生物群落变体是特定的,与宿主的总体形态结构和生活方式无关。不同类群宿主为共生桡足类动物提供的五种微生物群落类型已被确定(表2)。1. 底栖无脊椎动物的体表作为微生物群落,其条件与其他任何基质上的外部环境相比,对桡足类动物的任何物理和化学影响几乎没有差异。显然,在这种情况下,营养依赖性起着决定性作用。适应发展存在某些方向,这在某种程度上是所有水生外寄生甲壳类动物的特征,并且具有一个功能任务——帮助外寄生虫附着在宿主体表。首先,小颚和颚足显著发育,它们形成大爪子的形状,桡足类动物用其牢固地附着在宿主体表,并能够在上面移动而不会有被冲走的危险。其次,身体形态经历背腹向的表达和前体的扩展,形成头盾,使共生体能够紧紧地压在宿主身体表面并避免失去宿主(表2)。有时,一些外寄生虫会刺激宿主皮肤组织形成瘿,这也为寄生虫提供了恒定的条件,而不会有失去宿主的任何威胁。然而,这种现象分布并不广泛,仅在某些甲壳类和棘皮动物宿主群体中观察到。2. 宿主生物体内的狭窄管状腔,它们要么是外部环境的一部分(如海绵的管道系统),要么是生物体内部环境的一部分(如血液系统的通道或消化系统的细小部分),其大小和形状总是受到严格限制。占据这种微生物群落的所有寄生虫的特征是强烈的形态变化。它们表现为腿部或任何其他附肢的退化(通常程度显著),在一定程度上失去分节,并呈现出毛虫状(或蠕虫状)的身体形态(表2)。这种微生物群落仅在一种情况下被外寄生虫占据(来自海绵管道系统的Spongicola uncifer),在所有其他情况下被内寄生虫占据。3. 与外部环境相连的大腔。各种起源的结构,如软体动物的外套腔、甲壳类动物的鳃腔和育儿袋、蛇尾类动物的囊腔和海鞘的鳃腔,都属于这种微生物群落类型。它们都具有与外部环境相对不同且体积较大(与桡足类动物的大小相比)的特点,这为寄生虫提供了免受外部环境因素影响的充分保护以及持续的食物来源,如宿主体内的成分或水流带来的食物颗粒。微生物群落栖息者的形态变化有两个方向。生活在双壳类动物外套腔中的绝大多数桡足类动物实际上没有形态变化。另一方面,甲壳类动物鳃腔和育儿袋、蛇尾类动物囊腔以及海鞘鳃腔的栖息者具有强烈的形态变化特征。通常表现为在一定程度上失去分节、附肢退化和身体肿胀,如Sphaeronella属的物种(表2)。在生命周期中也观察到变化:发育阶段有减少的趋势(无节幼体阶段在卵巢覆盖下发育)。在这种情况下,桡足幼体阶段从卵中孵化出来。4. 宿主生物体内的腔。不同类群宿主中的这种微生物群落类型有不同程度的体现。我们在多毛类动物的体腔、软体动物的腔隙系统、甲壳类动物的混合体腔、棘皮动物的体腔以及海鞘的身体腔中都能识别到它。占据这种微生物群落的桡足类动物有两个基本的进化方向。第一种情况,寄生虫不发生形态变化,保持其结构的原始模式,如同祖先的自由生活形式。第二种情况,寄生虫发生强烈的形态变化,它们要么直接生活在腔液中,要么被包囊包围(如Cucumaricolidae科)。5. 最后一种类型的微生物群落最为特殊。在外部环境(二级环境)和内部环境(一级环境)两种环境中同时存在,导致寄生虫在结构和组织水平上完全失去祖先类型。同时,寄生虫身体在形态和功能上分为两部分,产生了一种新的结构——外质体和内质体。在这种情况下,我们涉及到中寄生现象。