Physiological adaptations of the living agnathans

Abstract

ABSTRACTThe modes of life and environments of the extant agnathans (cyclostomes) are discussed in relation to their adaptations to temperature, light, oxygen and salinity. As their antitropical distribution indicates, both hagfishes and lampreys are cold water groups. Since hagfishes live in deeper waters than lampreys, they are not exposed to the marked seasonal changes in temperature and light which influence major events in the lamprey life cycle. Both groups tend to be nocturnally active, either burrowing during daylight as in the case of larval lampreys (ammocoetes) and most hagfishes, or showing cryptic behaviour as in the case of adult lampreys. Olfaction plays a major part in the location of prey, presumably aided in adult lampreys by their eyes and sensitive electrosensory system. Rates of standard oxygen consumption, ventilatory frequency and heart rate of adult lampreys increase at night. Standard oxygen consumption is relatively low in ammocoetes (as it also is in hagfishes) but increases markedly during metamorphosis into the adult lamprey. Ammocoetes and hagfishes, and to a lesser extent adult lampreys, are resistant to reduced environmental oxygen tensions. Differences in the oxygen dissociation curves of ammocoetes, adult lampreys and hagfishes can be related to differences in the characteristics of their monomeric haemoglobins and their environments and modes of life. The extraordinary tolerance of the hagfish heart to hypoxia is a reflection of a robust capacity for glycolysis, an LDH isozyme geared towards anaerobic functioning and a low work output. The hagfishes, which are restricted to marine waters, are osmoconformers. The osmolality of their blood, which is almost wholly attributable to inorganic ions, is virtually identical to that of full strength sea water (c. 1000 mOsmkg−1). By contrast, the osmolality of the blood of larval and adult lampreys when in fresh water is only 205-260 mOsm kg−1, i.e. about one quarter to one fifth of those of hagfish, and these rise only to 240-270 mOsm kg−1in the adults of anadromous lampreys in sea water. The regulation of ions by adult lampreys is achieved by mechanisms similar to those adopted by teleosts. The implications of the contrasting ionic and osmotic physiology of the two living groups of agnathans are discussed in relation to their possible environmental history and against the background of their Carboniferous fossil representatives.