Life in the Open Ocean. Joseph J. Torres

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СКАЧАТЬ of motor control (Hessinger 1988). Because mice are not within the normal prey spectrum of jellies, fiddler crabs have been used as test subjects for effects of cnidarian venom. Their symptoms include violent motor activity, paralysis, and in some cases, autotomy (spontaneous loss) of the walking legs (Hessinger 1988).

      Most of the toxic effects associated with cnidarian venoms can be traced to compromised cell membranes. Excitable tissues such as nerve and muscle are particularly susceptible to membrane disruption because they require an intact membrane to function. Cardiovascular distress, loss of motor control, and violent convulsions are all indicative of compromised cell membranes in the neuromuscular system and will nearly always be the most obvious symptoms obtained when an experimental subject is injected with a purified venom.

      That said, a world of difference exists between a jellyfish sting and the impacts of purified jellyfish venom, or our beaches would be empty much of the time. Only a very few people die each year from jellyfish stings, and many of those fatalities are due to an allergic response like that seen with bee stings. Most people only feel a minor bit of irritation that goes away in a few hours, even with a venom that is as potent as that of a rattlesnake. Why? The amount delivered by a jellyfish sting is minute compared with the amount delivered by a snake. A large rattler can inject volumes of a milliliter or more of pure venom when it strikes, which makes it very deadly. The larger jellies such as the Man O’ War and the Australian sea wasp can deliver large enough quantities of venom to cause severe distress or worse. For small creatures such as fish larvae and many species of zooplankton though, the amount of venom introduced by small jellyfish stings is enough to disable them. The many small “harpoons” of the nematocysts trap them on the tentacle, and they can then be conveyed to the mouth.

      Interaction with Prey

As noted above, medusae are radially symmetrical tentaculate predators that rely on contact to entangle their prey. They are not “sight‐hunters” or raptors (e.g. Greene 1985) capable of actively seeking a prey item and chasing it down. Yet despite their seeming inability to select prey, virtually all gelatinous predators for which electivity indices have been calculated show some degree of selectivity in their diet (Purcell 1997).

      Most pelagic cnidarians feed mainly on copepods, the dominant metazoan zooplankton group in the vast majority of the world ocean (Purcell 1997). This is not surprising; a nonliving bit of “marine flypaper” cut into the shape of a medusa and floating with the ocean current would mainly snag copepods. It is the exceptions to the rule that are intriguing and lead one to wonder about how the selectivity is achieved. For example, Purcell (1997) observed that the hydromedusae Bougainvillia principis and Proboscidactyla fed mainly on barnacle larvae and molluscan veliger larvae, respectively. Are cnidarian dietary preferences the result of a limited prey field or actual selectivity? We will investigate cnidarian hunting from a theoretical perspective. Interactions with prey have two basic elements: the encounter and the capture (Purcell 1997). Factors influencing the encounter phase have been treated in a number of studies and may be divided into four subcategories: direct interception, encounter zone, water flow and swimming, and attraction between predator and prey.

      Direct Interception

Cnidarian tentacles may be considered as a large, loosely configured filter, and the concept of direct interception derives from filtration theory (Rubenstein and Koehl 1977; Purcell 1997). Because the spacing of tentacles in any cnidarian predator is usually much greater than the prey diameter, particularly for small prey, the direct interception of a prey item on a tentacle depends only on the diameter and swimming speed of the prey and the diameter of the tentacle. Further, the theory predicts that larger, faster prey would be selected for by tentaculate predators generally. It applies most directly to ambush predators such as the siphonophores and some of the medusae. Figure 3.16 shows how the ubiquitous medusan predator Pelagia noctiluca captures prey on one of its tentacles while swimming. Prey is trapped by nematocyst discharge and conveyed to the mouth with the cooperation of the oral arms as the Pelagia continues to swim.

      Encounter Zone

Madin (1988) used field observations, videography, and measurements to produce a very useful conceptual model of feeding behavior in medusae, ctenophores, and siphonophores. In his model, the shape of the medusa’s bell, its arrangement of tentacles, and its swimming behavior create an encounter zone where probability of prey capture is maximized. Within the encounter zone, the likelihood that various prey types will be caught depends on the interaction of tentacle density, tentacle spacing, prey size, type, and the behavior and properties of nematocysts (Table 3.1).

      Figure 3.17 illustrates modes of tentacle deployment for a variety of different medusae and siphonophores. Most of the medusae illustrated are hydromedusae, underscoring the diversity in their morphology. “Type” species represented in Figure 3.17 are listed in Table 3.1 and described briefly below.

       Anthomedusae

      Calycopsis typa (Figure 3.17a) has a globular bell and thick tentacles held out radially when fishing, creating a discoid volume about three times the diameter of the bell.

Stomotoca pterophylla (Figure 3.17b) is a species with two tentacles that may be 50 times the bell diameter when fully extended. The tentacles occupy only a tiny fraction of the spherical volume accessible to them. Prey must swim into the tentacles for successful capture. Stomotoca feeds on medusae that are large enough to allow its feeding apparatus to function effectively.

Figure 3.16 Feeding behavior of Pelagia noctiluca on motile prey. (a) When the prey touches a marginal tentacle, there is an immediate nematocyst discharge, followed by a tentacle contraction after 2–3 seconds; (b) the stiff tentacle bends toward the nearest oral arm, at the same time the oral arm moves upward, turns slightly, and draws its endodermic layer near the food; (c) the stiff tentacle releases the prey and moves upward, going away from the oral arm; (d) the oral arm grasps the prey completely and starts the peristaltic and mucous movements, which drive the food to the oral arm groove, to the manubrium, and finally to the gastric cavity. The inset figure is a transverse section of “d.”

      Source: Rottini‐Sandrini and Avian (1989), figure 1 (p. 52). Reprinted by permission from Springer Nature Customer Service Centre GmbH Springer‐Verlag, Marine Biology, Feeding mechanism of Pelagia noctiluca (Scyphozoa: Semaeostomeae), Rottini‐Sandrini and Avian, 1989.

Table 3.1 Tentacular encounter zones and deployment patterns, tentacle length, volume, and density, and prey types of some medusae and siphonophores.

      Source: Adapted from Madin (1988), table 2 (p. 423).

Species	Encounter zone – shape of space	Tentacle deployment a pattern (see СКАЧАТЬ В начало < 52 53 54 55 56 57 58 59 60 61 > В конец