Respiration and Circulation in Land Crabs: Novel Variations on the Marine Design

Both the "true" crabs (Brachyura) and hermit crabs (Anomura)include species that show numerous behavioral, morphological,and physiological specializations permitting terrestrial life.This paper examines respiratory and circulatory adaptationsfor air breathing in these land crabs. Respiratory specializationsinclude modification of gas exchange structures for air breathing(gills and elaborated branchial chamber linings), ventilatorymechanisms permitting effective air pumping, an elevated hemolymphoxygen capacity, and a primarily CO₂- rather than O₂- sensitiveventilatory control system. The qualitative aspects of hemolymphoxygen transport and metabolic rate are apparently unchangedfrom that of marine crabs. While the basic cardiovascular morphologyof land crabs appears similar to that of marine forms, thereis considerable elaboration of the vasculature of the branchialchamber lining, which in some species includes a unique doubleportal system. Cardiac output is lower in land crabs (probablyrelated to their higher hemolymph O₂ capacity), but insufficientdata on hemolymph pressures prevent comparisons with marineforms. In general, land crabs have modified (sometimes extensively)existing structures and processes found in their marine relativesrather than evolving structures for terrestrial life de novo.Accordingly, land crabs present a useful model for the evolutionof terrestriality, showing that even subtle anatomical changescan result in the large changes in physiological function necessaryfor the terrestrial invasion.     

Form and Function in Reptilian Circulations

Consistent with the great variation in their circulatory morphology,there are distinct variations in the cardiovascular physiologyof extant reptiles. The chelonian and squamate reptiles havea complexly structured heart that includes three partially separatedventricular cava. In most species (under most conditions), theventricle acts as a single pressure pump perfusing both thepulmonary and systemic circuits. However, the varanid lizardsprovide a striking exception. Subtle evolutionary changes incardiac morphology allow the ventricle of the varanid lizardto divide functionally during systole into a low pressure, pulmonarypump and a high pressure, systemic pump. The crocodilians representyet another anatomical and physiological pattern. The ventricleis completely divided into left and right chambers as in homeotherms,but the systemic and pulmonary circuits may still communicatethrough the left aorta that arises from the right ventricle. A fundamental feature of all reptilian circulations is the abilityto regulate the distribution of cardiac output between systemicand pulmonary circuits via central vascular blood shunts.Regardlessof species, mechanisms for regulating intracardiac shuntinginvolve changes in the balance between peripheral resistanceof the pulmonary and systemic circulations, and adjustmentsin cardiac performance per se. Several hypotheses are presentedthat suggest selective advantages for central vascular shuntingin intermittent breathing reptiles with variable body temperatureand metabolic rate.     

CUTANEOUS GAS EXCHANGE IN VERTEBRATES: DESIGN, PATTERNS, CONTROL AND IMPLICATIONS

1. The exchange of oxygen and carbon dioxide between skin and environment is commonplace in the vertebrates. In many lower vertebrates, the skin is the major or even sole avenue for respiration.
2. As implied by the physical laws governing diffusion of gases, the skin diffusion coefficient, surface area, gas diffusion distance and transcutaneous gas partial pressures may independently or jointly affect cutaneous respiration. In vertebrates, each of these variables has undergone modification that may be related to dependence upon cutaneous gas exchange.
3. Both theoretical models and experimental data suggest that cutaneous gas exchange is limited by the rate of diffusion. However, changes in convection of the respiratory medium and of blood may partially compensate for diffusion limitation, and potentially function in the regulation of cutaneous gas exchange.
4. Typically, the skin is one of several gas exchangers, although many salamanders and some species in other vertebrate groups breathe solely through the skin. The cutaneous contribution to overall gas exchange is often most important in small animals, at cool temperatures, at low levels of activity and in normoxic and normocapnic conditions. Branchial and pulmonary respiration increasingly predominate in other circumstances.
5. Often, the skin figures more prominently in CO₂, excretion than in O₂, uptake.
6. Cutaneous gas exchange emerges in vertebrates as a process perhaps less effective and more constrained than branchial or pulmonary exchange but also less energetically costly. Its utility is indicated by its wide and successful exploitation in vertebrates occupying a diverse array of habitats.     

Identifying and Evaluating Patterns in Cardiorespiratory Physiology

SYNOPSIS. A major theme of many papers in this symposium isthe identification of broad physiological trends and patternsthat extend beyond the boundaries of data from individual studies.Recognizing patterns in everything from hypoxic ventilatorypatterns to regulation of blood gases not only helps the investigatorunderstand specific data sets, but also helps place those datain a broad context. Yet, recognizing physiological patternsis confounded by two factors: phylogenetic relationships andphysiological state. Fortunately, the last decade has seen infiltrationof sound evolutionary theory, including tools of cladisiticanalysis and population genetics, into more and more studiesof comparative physiology. However, even when an experimentercarefully accounts for phytogeny, differences in physiologicalstate in the experimental animals can still obscure physiologicalpatterns. Two informal categories of physiological state aredescribed, the first obvious and frequently controlled for,and the second less obvious and typically not controlled for.Examples of the latter, including seasons, rhythms, prandialeffects and sex of the animal, are developed to show how ignoringthese can lead to considerable misleading variation in cardiorespiratorydata sets. Considering physiological state is vital in producingreliable data that can be used meaningfully for delineatingbroad physiological patterns.     

The Respiratory Transition from Water to Air Breathing During Amphibian Metamorphosis

SYNOPSIS. Profound developmental changes occur in the morphologyand physiology of the respiratory system of amphibians duringthe transition from strictly aquatic to dual aquatic-aerialbreathing. This developmental transition usually involves modificationsin ventilatory mechanisms and/or respiratory surfaces {e.g.,degeneration of gills, ventilation of functional lungs). Boththe first appearance of obligate air breathing and the subsequentdependence upon it by amphibian larvae are sensitive to a varietyof environmental stressors during critical developmental periods.These stressors include oxygen availability, ambient temperature,the risk of predation and mode of feeding.     

Role of the Central Circulation in Regulation of Cutaneous Gas Exchange

SYNOPSIS. Although primarily limited by the rate of diffusionof oxygen and carbon dioxide across the integument, cutaneousgas exchange is also affected by adjustments in the absoluteflow of blood to the skin, the pattern of distribution of bloodwithin the cutaneous vascular bed, and the effects of centralvascular shunting on gas partial pressures of arterial bloodperfusing the skin. The interplay of these various factors,particularly in animals with heterogeneous arterial supply tothe skin and/or with highly variable intracardiac shunts, potentiallyis complex, only poorly understood, and worthy of considerablefuture experimentation.