The body plan of the cnidarian medusa: distinct differences in positional origins of polyp tentacles and medusa tentacles
The body plan of the cnidarian medusa: distinct differences in positional origins of polyp tentacles and medusa tentacles
复制标题
刺胞动物水母的身体结构:水螅触手和水母触手位置起源的明显差异
作者:
H. Shimizu;H. Namikawa
Cnidarians have two typical body forms, the attached form (polyp) and the swimming form (medusa) (Fig. 1). The polyp has a mouth with tentacles closely surrounding it at the apical end of a cylindrical body column (Fig. 1A). The morphology of a medusa is characterized by a body like an inverted bowl, termed the umbrella (Fig. 1B). The umbrella has tentacles hanging down from its margin (marginal tentacles). The mouth is located at the end of a columnar structure termed the manubrium, which is located at the center of the umbrella. Despite these significant differences in morphology, zoology textbooks typically state that ‘‘the medusa is the upside down form of a polyp,’’ implying that the two forms have basically the same body plan (Ruppert and Barnes 1996). As a result, it has generally been assumed that the tentacles of the polyp and the marginal tentacles of the medusa are homologous. However, it remains unknown whether the textbooks’ description is correct or not. Based upon classical observations and recent results of molecular studies, we present a simple but novel answer to this question by showing that animals that belong to Cnidaria have two areas in the body where tentacles are formed, one at the oral end, and the other in an aboral location on the animal. We focused our analysis on the differences in positional origins of hydropolyp tentacles and hydromedusa marginal tentacles in this study. Although the medusa form occurs in three cnidarian classes (viz., Scyphozoa, Cubozoa, and Hydrozoa), of these about 75% of the species that have the medusa stage belong to Hydrozoa (Brusca and Brusca 1990; Boero et al. 1992; Bouillon and Boero 2000). Furthermore, Hydrozoa is currently the only class where molecular analysis has been performed, although extensive gene expression studies have been carried out mostly with polyps. The term tentacle is generally used to describe the flexible appendages that are located near the mouth opening in cnidarians and are involved in capturing prey using nematocytes (Ewer 1947). If a tentacle produced in the polyp showed developmental continuity into the medusa, this would be direct evidence that the polyp tentacles and the medusa tentacles are essentially the same structure. This continuity, however, is not seen. Instead, medusa tentacles are newly formed during the process of medusa formation (namely, asexual budding in Hydrozoa). The morphology ofHydra, a hydrozoan research model, is characterized by the solitary polyp form with the hypostome surrounded by tentacles at the oral end, a large body column/ gastric region, and with the peduncle and foot region at the aboral end (Fig. 1C). Hydra has been considered to represent the typical solitary polyp form of this class. However, some hydrozoans have polyps in which tentacles emerge in two regions. A polyp of the hydrozoan family Corymorphidae (Fig. 1D) has tentacles that are at a glance concentrated at the oral end, suggesting that the morphology of the animal is basically similar to Hydra. This view is, however, entirely erroneous. The digestive region (denoted by DR) of a hydra polyp, which is pink in color because of digested food content, occupies about 3/4 of the body column length, whereas the rest (about 1/4) corresponds to the peduncle region (Fig. 1C). In contrast, the digestive region of a corymorphid polyp is the region that lies between the two tentacle rings occupying only about 1/10 of the body. The rest of the polyp (about 9/10) is occupied by the peduncle region (Fig. 1D), demonstrating that the two polyps have different proportions of the body parts in apparently similar cylindrical morphology. Therefore, the position of the secondary tentacle ring of the corymorphid polyp corresponds, in a hydra polyp, to the boundary between the digestive region and the peduncle region, which is located on the aboral side of the animal. Because the secondary tentacle ring of the corymorphid polyp is located aboral to the digestive region of the animal, the tentacles in the secondary ring should be considered aboral tentacles. This implies that there are two tentacle regions in the corymorphid polyp, one on the oral side of the digestive region and the other in the aboral side of this region. It should be noted that the tentacles in the two regions are involved in capturing prey in a cooperative manner. When three subclasses of Hydrozoa (Limnomedusae, Anthmedusae [Athecata], and Leptomedusae [Thecata]) are EVOLUTION & DEVELOPMENT 11:6, 619 –621 (2009)
DOI:
10.1242/dev.122.12.3785
发表时间:
1996
期刊:
Development (Cambridge, England)
影响因子:
--
作者:
Millet,S;Bloch-Gallego,E;Simeone,A;Alvarado-Mallart,RM
通讯作者:
Alvarado-Mallart,RM