SUMMARY In hearts of higher invertebrates as well as vertebrates, the work performed by the ventricle is a function of both rate and contractility. Decapod crustaceans experience a hypoxia-induced bradycardia that is thought to result in an overall reduction in cardiac work; however, this hypothesis has not yet been tested and is the primary purpose of this study. In the grass shrimp Palaemonetes pugio, cardiac pressure and area data were obtained simultaneously, and in vivo, under normoxic (20.2kPaO2) and hypoxic (6.8 or 2.2kPaO2) conditions and integrated to generate pressure 阂area (P 阂A) loops. The area enclosed by the P 阂A loop provides a measure of stroke work and, when multiplied by the heart rate, provides an estimate of both cardiac work and myocardial O2 consumption. Changes in intra-cardiac pressure (dp/dt) are correlated to the isovolemic contraction phase and provide an indication of stroke work. At both levels of hypoxic exposure, intra-cardiac pressure, dp/dt, stroke work and cardiac work fell significantly. The significant decrease in intra-cardiac pressure provides the primary mechanism for the decrease in stroke work, and, when coupled with the hypoxia-induced bradycardia, it contributes to an overall fall in cardiac work. Compared with normoxic P 阂A loops, hypoxic P 阂A loops (at both levels of hypoxia) become curvilinear, indicating a fall in peripheral resistance (which might account for the reduction in intra-cardiac pressure), which would reduce both stroke work and cardiac work and ultimately would serve to reduce myocardial O2 consumption. This is the most direct evidence to date indicating that the hypoxia-induced bradycardia observed in many decapod crustaceans reduces cardiac work and is therefore energetically favorable during acute exposure to conditions of low oxygen.
ABSTRACT The passage of a barium meal (15 % by mass) was followed through the digestive system of the blue crab Callinectes sapidus by flash-freezing crabs at set intervals, followed by radiography of specimens. Food moved from the oesophagus into the stomach region within 15 min. After 1–2 h, food was visible in the midgut, at 6 h it had reached the hindgut, and material was still present in the stomach at this time. The stomach was emptied between 8 and 10 h after feeding, and the entire digestive system was cleared of material after 18 h. A pulsed-Doppler flowmeter was used to monitor cardiac variables and arterial haemolymph flows during a 4 h control and 24 h postprandial period. Heart rate increased immediately upon food detection and remained elevated for 16–18 h after food ingestion. There was no significant change in stroke volume of the heart, and total cardiac output increased significantly and remained elevated above pre-feeding levels for 24 h after feeding. There was no change in haemolymph flow through the anterior or posterior aorta, but flow increased in the sternal, anterolateral and hepatic arteries. These changes in haemolymph flow reflected the use of the chelae and mouthparts in feeding, contraction of the visceral muscle surrounding the gut system and mobilisation of enzymes from the hepatopancreas. There was also a postprandial increase in the rate of oxygen uptake (apparent specific dynamic action). The rate of oxygen consumption reached maximal levels 4 h after feeding and decreased slowly thereafter, reflecting the increased use of oxygen in digestion and absorption.
Hemoglobin (Hb) is a highly conserved protein that provides a vital link between environmental oxygen and its use and/or storage within an organism. While ubiquitous among vertebrates, Hb occurs frequently in invertebrate phyla as well. Many arthropod species use the copper-binding pigment hemocyanin, but unique in this phylum are the branchiopod crustaceans, which express Hb. Branchiopod Hb concentration and structure are exquisitely sensitive to environmental oxygen availability. Hemoglobin concentration and oxygen-binding affinity increase with decreasing oxygen tension in Daphnia, Artemia and Triops. The change in binding affinity is attributed to differential Hb subunit expression in Daphnia and Artemia but remains unclear for Triops. This is the first study to demonstrate developmental plasticity of Hb subunit expression in a notostracan, Triops longicaudatus, reared under conditions of varying oxygen availability. In response to variable oxygen environments, T. longicaudatus differentially express four primary Hb subunits ranging between 30 and 34 kDa, with normoxic-reared animals expressing primarily the heavier subunits, and hypoxic-reared animals expressing increased proportions of the lower molecular mass subunits. Moreover, differential Hb subunit expression is induced upon transfer of normoxic-reared adults to a hypoxic environment, such that the distribution of Hb subunits in the transferred adults becomes similar to that of hypoxic-reared animals. Two-dimensional gel electrophoresis and follow-up analyses revealed several isoforms of Hb subunits that may represent differential gene expression and/or post-translational modification. Unlike Daphnia and Artemia, the Hb hypoxic response in Triops is not reversible in that there was no significant decrease in Hb concentration or change in Hb subunit expression pattern when hypoxic-reared adults were transferred to a normoxic environment.
Summary In many crustaceans, female reproduction represents a time of increased metabolic demand. Palaemonetes pugio are typically hypoxia tolerant; but the energetic demands of reproduction may compromise their ability to tolerate hypoxic conditions. Given the correlation between cardiac output and metabolic demand, we used cardiac output (CO) to measure differences in metabolic demand in the life‐history stages of P. pugeo . We hypothesized that (1) the cost of egg production would result in an increased CO for gravid females compared with non‐gravid females; (2) those females that were both ovigerous and gravid would have an additional metabolic demand due to brooding behaviour (pleopod fanning) and hence an even greater CO; and (3) hypoxia tolerance would decrease with increasing reproductive demand. To test these hypotheses, we compared cardiac output across three reproductive states and at decreasing water oxygen tensions. Ovigerous females had significantly greater pleopod fanning frequency than non‐ovigerous females at all oxygen tensions. Additionally, ovigerous/gravid females had significantly higher cardiac output at all oxygen tensions than gravid only or non‐gravid females. Changes in cardiac output indicate that females became more sensitive to environmental oxygen tension with increasing reproductive demand. Non‐gravid females were able to maintain cardiac output down to 15 mm Hg O 2 , whereas gravid and ovigerous/gravid females maintained cardiac output down to 50 mm Hg and 75 mm Hg O 2 , respectively. These differences in CO suggest that metabolic demands of females change with reproductive state and, while gravid and ovigerous/gravid females appear more sensitive to low oxygen tensions, they are able to physiologically tolerate low environmental oxygen conditions.
ABSTRACT Oviparous amniotes exchange respiratory gases across a specialized vascular membrane, the chorioallantois. Although many investigations into the physiology of amniote eggs, particularly those of the chicken, have been carried out, there is no comparative information about the chorioallantoic membrane. Given the differences in size and phylogeny, it might be expected that significant morphological differences exist in this gas-exchange organ.