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Exoskeleton, Distribution, and Movement of the Flexible Setules on the Myodocopine (Ostracoda: Myodocopa) First Antenna

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Exoskeleton, Distribution, and Movement of the Flexible Setules on the Myodocopine (Ostracoda: Myodocopa) First Antenna EXOSKELETON, DISTRIBUTION, AND MOVEMENT OF THE FLEXIBLE SETULES ON THE MYODOCOPINE (OSTRACODA: MYODOCOPA) FIRST ANTENNA Andrew R. Parker ABSTRACT The halophore, halothalium, and s-seta are termed herein and occur on the myodocopine first an- tenna, probably the most systematically significant myodocopine appendage. The morphology of Downloaded from https://academic.oup.com/jcb/article/18/1/95/2418948 by guest on 29 September 2021 the exoskeleton and distribution of these structures are described, following scanning electron and light microscopic analyses and literature comparisons. The movements of these and other myo- docopine first antennal structures are studied, using video recordings of an exemplary species. Halo- phores are setules with a characteristic exoskeletal ultrastructure comprising a layer of very fine rings, with walls approximately circular in cross section, covered by an outer, probably elastic, layer or sheath. There is a single pore at the halophore tip. This organization permits great flexi- bility. If dendrites innervate halophores, the flexibility may aid in sampling for water-borne chem- icals or mechanoreception. The s-seta (formerly the sensory seta) is a seta arising from the fifth ar- ticle of the myodocopine first antenna that frequently possesses numerous long halophores. The s- seta is always and only present in the Myodocopina. The collective halophores distributed over the whole of one first antenna are termed the halothalium. Analogies of the structures described in this study are made with other crustacean structures. The ostracod suborder Myodocopina, of at least some of the first antennal setae was within the order Myodocopida, comprises five believed to be tactile (Vannier and Abe, 1993). extant families; Cypridinidae, Cylindrole- In addition, the function of the seta arising berididae, Philomedidae, Rutidermatidae, and from the fifth article has long been designated Sarsiellidae (Kornicker, 1975). Myodoco- as sensory (e.g., Skogsberg, 1920; Sars, 1922). pines occur in marine or brackish environ- However, no such evidence has been pre- ments world-wide at all depths (Cohen, sented to substantiate these sensory claims. 1982). Most (including juveniles) can swim, Presumably, based on morphological simi- but are benthic or epibenthic (only 3-6 of the larities, original assumptions were made from 26 described cypridinid genera are wholly or comparisons with homologous or convergent mostly planktonic) for much of their lives structures in other crustacean taxa. Poulsen (Cohen, 1982). They all exhibit sexual di- (1962) followed the terminology for myo- morphism (Cohen and Morin, 1990). They docopine limbs as established by previous are "filter-feeders" (Kornicker, 1975), or more myodocopine workers, but remained uncon- accurately comb-feeders, because they typi- vinced of the implication about their function. cally operate at low Reynolds numbers (Co- hen, 1989), scavengers, active predators, "col- Morphology of the Myodocopine First An- lectors" (collect detritus from the sediment; tenna.-The myodocopine first antenna con- Walker, 1972), detritus feeders (Cannon, sists of eight articles (some occasionally 1933; Kornicker, 1975), or parasites (Bennett fused). The more proximal articles bear var- et al., 1997). The first antenna is probably the ious (often short) setae, and three of the four most significant appendage in myodocopine distal articles (namely 5, 7, and 8) bear about systematics. seven relatively long setae. It is these longer setae that project through the anterior end of Previous Terminology.-The first antenna the open carapace and probably reach out be- (antennule) of Myodocopina includes about yond the boundary layer of flow (ostracods seven long terminal and subterminal setae of typically operate only at low Reynolds num- previously unconfirmed function (Fig. 1 ). The bers; see Koehl and Strickler, 1981). These first antenna of vargula hilgendorfzi Poulsen setae are historically termed: the sensory seta (Cypridinidae) extended when the animal was (arising from the fifth article); the a- (usu- resting on the substrate. Thus, the function ally short), b- (usually mid-long) and c-setae Downloaded from https://academic.oup.com/jcb/article/18/1/95/2418948 by guest on 29 September 2021 Fig. I. Azygocypridina lowryi, adult female, left first antenna, medial view (eighth article not in view). New termi- nology (for explanation see discussion section) is used in the labeling; traditional terminology of s-seta is within parentheses. (from the seventh article); and the d-, e-, f-, and glion to the muscles, but other antennulary g-setae (from the eighth article) (Fig. 1). Some nerve fibers terminate in cells in the postero- setae are missing in a few myodocopine taxa. medial corner of the ganglion. From here, Most are annulate, and may bear spines, se- other fibers extend to the distal end of the tules, or "suckers" (the last in males only). The ganglion, passing out as a bundle running to sensory seta of the fifth article usually appears the tip of the limb. Between these two bun- morphologically distinct. This seta of medium dles of nerves lies a group of four or five length is sometimes widened (or bulbous) at bipolar cells, giving rise to medium-sized gi- the base or has a separated basal section, and ant fibers (Cannon, 1931). The antennulary usually has at least six long thin flexible se- nerves are similar in the cypridinid Giganto- tules. These setules, previously unconvention- cypris mulleri Skogsberg (see Cannon, 1940). ally termed filaments (see Watling, 1989) have However, nothing more detailed regarding the been described as unringed, although in the lit- precise innervation of the first antennal setae erature they are often illustrated with stippling, has been published. giving them a granular appearance (e.g., Poulsen, 1962). This is how they appear at Aims.-The aims of this study are to: (1) de- magnifications less than about 250x. scribe the morphology of the halophore ex- This study demonstrates that the setules of oskeleton using the scanning electron micro- the b- to g- and the sensory first antennal se- scope (SEM), (2) describe the distributions of tae of Myodocopina share a characteristic ul- halophores within myodocopine taxa using trastructure and all are termed "halophores" the literature and light microscopy, (3) de- herein. The sensory seta is termed the "s-seta." scribe the movement of the myodocopine first The collection of halophores on one first an- antennal setae and setules using video record- tenna is termed the "halothalium." The terms ings, (4) compare the flexible setules of the halophore, s-seta, and halothalium, designated myodocopine first antenna with other crus- in this paper, are used hereafter to avoid un- tacean structures, and (5) review the litera- necessary confusion (an explanation of these ture on, and standardize the terminology of, terms can be found in the discussion section). myodocopine first antennal setae/setules. Techniques suitable for the study of nervous Myodocopine First Antennal Nervous Sys- structures are not employed. tem.-The nervous supply of the first antenna of the cypridinid Doloria levis Skogsberg MATERIALS AND METHODS arises from two swellings on the ventral side Specimens Examined.-Living specimens of Azygo- of the forepart of the nerve ring, marking the cypridina lowryi Kornicker were collected in single- deuterocerebral part of the brain (Cannon, chamber baited traps designed by Keable (1995). These 1931). The antennulary basal ganglion is the were set overnight at depths of 200 m (34°31.48'S, 151°13.22'E) and 300 m (34°31.80'S, 151°15.60'E) off largest in the body of D. levis and occupies the Wollongong (New South Wales, Australia) coast. The most of the basal article. What appear to be ostracods were removed from the traps and immediately motor nerves pass directly through the gan- transported to the laboratory in fresh aerated sea water. Table 1. Material examined (museum specimens fixed in 7% Formalin solution and preserved in 70% ethanol). AM represents Australian Museum (Sydney); NMNH represents National Museum of Natural History (Washington, D.C.). Downloaded from https://academic.oup.com/jcb/article/18/1/95/2418948 by guest on 29 September 2021 Behavioral observations were carried out within 4 h, dur- ing which time the ostracods appeared to be in a healthy condition. Preserved museum specimens (fixed in 7% Formalin solution and preserved in 70% ethanol) of ostracods and other crustaceans were examined (Table 1). Anatomical Study.-For SEM examination, preserved mu- seum specimens (Table 1 ) were cleaned, using 5 half-sec- ond exposures to ultrasound, critical-point dried, using a BioRad CPD 750, then coated with gold. These speci- mens were examined using a Cambridge Instruments S 120 SEM. Fig. 2. Diagrammatic sections of the halophore exo- Accidentally broken halophores of A. lowryi were stud- skeleton. A, longitudinal section (shaded areas represent Downloaded from https://academic.oup.com/jcb/article/18/1/95/2418948 by guest on 29 September 2021 ied to view the internal surface of their exoskeletons. In cross sections of the walls of the rings); B, cross section some cases, the rings which comprise the exoskeleton of through t-t. the halophores were themselves broken, and their inter- nal architecture was opportunely examined. The setae and setules of myodocopine limbs other than the first antennae were also examined under the SEM to The bases of the halophores are not finely an- search for external morphologies similar to those of halo- nulate, except where they arise terminally on phores. Azygocypridina lowryi was mainly used in this a seta. Each halophore bears a thin outer layer investigation because of its relatively large size (carapace (sheath) (Figs. 2, 3F), which follows the length about 11 mm). Preserved museum specimens of grooved external contours of the halophore species from other ostracod taxa and crustacean classes (Table 1) were examined similarly. set by the rings. Some myodocopines bear first antennal setae (other than the b- to g- or Behavioral Study.-Video recordings were made of sev- s-setae) with unringed setules, although these eral living specimens ofA.
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