Morpho-functional characterization of the goldfish (Carassius auratus L.) heart

Using morphological and physiological approaches we provided, for the first time, a structural and functional characterization of Carassius auratus L. heart. Besides to the classical four chambers, i.e. sinus venosus, atrium, ventricle, bulbus, we described two distinct structures corresponding to the atrio-ventricular (AV) region and the conus arteriosus. The atrium is very large and highly trabeculated; the ventricle shows an outer compacta, vascularized by coronary vessels, and an inner spongiosa; the bulbus wall is characterized by a high elastin/collagen ratio, which makes it extremely compliant. Immunolocalization revealed a strong expression of activated "eNOS-like" isoforms both at coronary endothelium and, to a lesser extent, in the myocardiocytes and the endocardial endothelium (EE). The structural design of the heart appears to comply with its mechanical function. Using an in vitro working heart preparation, cardiac performance was evaluated at different filling and afterload pressures. The hearts were very sensitive to filling pressure increases. Maximum Stroke volume (SV=1.08 ± 0.09 mL/kg body mass) was obtained with an input pressure of 0.4 kPa. The heart was not able to sustain afterload increases, values higher than 1.5 kPa impairing its performance. These morpho-functional features are consistent with a volume pump mechanical performance.     

Heart morphology in wild and farmed Atlantic salmon Salmo salar and rainbow trout Oncorhynchus mykiss

The normal shape of the salmonid ventricle is a triangular pyramid with the apex pointing caudoventrally. A strong positive correlation has been established between this shape and optimum cardiac output and function. Domesticated salmonids appear to have developed a more rounded ventricle with misaligned bulbus arteriosus. Several reports from fish health veterinarians indicate that fish with abnormal heart morphology have a high mortality rate during stress-inducing situations like grading, transportation and bath treatments. The present paper compares and describes the ventricle morphology of wild vs. farmed Atlantic salmon, and wild steelhead (anadromous rainbow trout) vs. farmed rainbow trout. Several parameters were measured to provide numerical measurement of the differences in shape, i.e. height:width ratio and the angle between the longitudinal ventricular axis and the axis of the bulbus arteriosus. We conclude that the hearts of farmed fish are rounder than those in corresponding wild fish, and that the angle between the ventricular axis and the axis of the bulbus arteriosus is more acute in wild fish than in their farmed counterparts. Further studies are necessary to reveal the prevalence, functional significance and possible causes of these abnormal hearts.     

Functional Morphology of the Heart in Fishes

The systemic heart of fishes consists of four chambers in series,the sinus venosus, atrium, ventricle, and conus or bulbus. Valvesbetween the chambers and contraction of all chambers exceptthe bulbus maintain a unidirectional blood flow through theheart. The heart is composed of typical vertebrate cardiac muscle,although there may be minor differences in the distributionof spontaneously active cells, the rate and nature of spreadof excitatory waves, and the characteristics of resting andaction potentials between different fish and other vertebrates.Cholinergic fibers innervate the heart, except in hagfish whichhave aneural hearts. Fish hearts lack sympathetic innervation.The level of vagal tone varies considerably, and is affectedby many factors. In some fish the heart is essentially aneural(without vagal tone) during exercise and may resemble an isolatedmammalian ventricle with increased venous return causing increasedcardiac output. There are many mechanisms that could increasevenous return in exercising fish. rß-adrenergic receptorshave been located on the hearts of some fish, and changing levelsof catecholamines may play a role in regulating cardiac activity.Changes in cardiac output in fish are normally associated withlarge changes in stroke volume and small cha-nges in heart rate.     

Cardiac function and critical swimming speed of the winter flounder (Pleuronectes americanus) at two temperatures

Using Transonic flow probes and a uniquely designed swimming flume, we directly measured cardiac parameters (Q, cardiac output; SV, stroke volume; and fH, heart rate) in winter flounder (Pleuronectes americanus) before and during critical swim speed (Ucrit) tests at 4 and 10 degrees C. Resting Q, SV and fH averaged 9.8 ml min(-1) kg(-1), 0.5 ml kg(-1) (1.0 ml g ventricle(-1)) and 21 beats min(-1) at 4 degrees C and 15.5 ml min(-1) kg(-1), 0.5 ml kg(-1) (0.95 ml g ventricle(-1)) and 34 beats min(-1) at 10 degrees C (Q10 values of 2.13, 0.91 and 2.35, for Q, SV and fH, respectively). Cardiac output, SV and fH increased by approx. 170%, 70% and 60% at both temperatures during the Ucrit test. However, cardiac parameters generally reached near maximal levels almost immediately upon swimming and remained at these levels until Ucrit (0.65 +/- 0.06 bl s(-1) at 4 degrees C and 0.73 +/ -0.07 bl s(-1) at 10 degrees C). This rapid rise in cardiac function to near maximal levels did not appear to be the result of stress alone, as Q only fell slightly when flounder were swum for 75 min at < 0.4 bl s(-1), speeds at which they appeared to swim comfortably. Our results suggest that both Q and Ucrit have been significantly overestimated in flatfishes, and that "lift-off"/slow swimming is energetically expensive. Furthermore, they show that maximum and resting stroke volume (per g of ventricle) are extremely high in the flounder as compared with other teleosts.     

The pericardium facilitates pressure work in the eel heart

An in siru perfused eel (Anguilla dieffenbachü) Gray 1842) heart was used to investigate the role of the pericardium in cardiac function. Hearts with intact pericardia were compared with hearts in which the pericardia were either punctured or opened completely. Cardiac function was assessed at low and high adrenaline levels by determining: maximum cardiac output; maximum sustainable output pressure; power output under maximal filling and output pressures; and maximum power output. Puncturing a small hole in the pericardium equalizing ambient and intrapericardial pressures had little effect on cardiac function and performance. Opening the pericardium, thereby fully exposing the chambers of the heart, severely limited the heart's ability to do pressure work. This effect was reversed at high adrenaline concentrations. Flow related work, and maximum power output levels were maintained after opening the pericardium.     

Cardiac performance of an isolated eel heart: effects of hypoxia and responses to coronary artery perfusion.

The pericardial sac containing the heart was removed from large (2.7-6.3 kg) long-finned eels (Anguilla dieffenbachii). Coronary arteries were cannulated in preparation for perfusion with eel Ringer or red cell suspensions. The hearts were maintained by Ringer perfusion while the performance of the heart was assessed. Responses of the hearts to increases in filling pressure and output pressure were recorded. Maximum cardiac output was 22.3 +/- 1.4 ml/min/kg body mass (mean +/- 1 SEM; N = 9). The highest cardiac power output was measured at maximum cardiac output and was 3.39 +/- 0.32 mW/g ventricle mass (mean +/- 1 SEM; N = 9). Eel hearts could sustain output pressures near 80 cm H2O, but cardiac output was reduced and cardiac power output was 1.89 +/- 0.24 mW/g ventricular mass (mean +/- 1 SEM; N = 9). Maximum cardiac output decreased by 14% when hearts pumped hypoxic Ringer with a PO2 of 11.5 torr. At high input pressures concomitant with high output pressures (80 cm H2O), cardiac power output decreased by 38% upon exposure to hypoxic Ringer. Coronary perfusion of hypoxic hearts with red cell suspensions (mean hematocrit 10.4%) at a rate of 2% of control cardiac output (0.20 ml/min/kg body mass) had no effect on maximum cardiac output. However, coronary perfusion enabled hypoxic hearts to maintain cardiac output when output pressure was raised to 80 cm H2O. Under conditions of high input pressure and high output pressure, power output increased by 20% compared to hypoxic hearts without coronary perfusion.(ABSTRACT TRUNCATED AT 250 WORDS)     

NO modulation of myocardial performance in fish hearts

In the mammalian heart, intracardiac nitric oxide (NO) regulates in an autocrine-paracrine manner cardiac function in the beat-to-beat response (Starling's law of the heart), short-term response (phasic control, e.g. excitation-contraction coupling, responses to neurotransmitters and endocrines) and long-term response (tonic control by altering gene expression). This trio of NO temporal-dependent actions has a long evolutionary history, as we have documented in the prototypic vertebrate heart, the teleost heart. This heart shares a common structural and functional scenario with higher vertebrate hearts exhibiting, at the same time, differences in myoarchitecture (trabecular vs. compact type), blood supply (lacunary vs. vascular) and pumping performance (sensitivity to filling pressure), thus providing challenging opportunities for revealing aspects of unity and diversity of cardiac NO in vertebrates. Using in vitro working teleost heart preparations we have shown that, under basal conditions, NO through a cGMP-mediated mechanism modulates ventricular performance (negative inotropism) and remarkably increases the sensitivity to filling pressure (i.e. the Frank-Starling response). NO-cGMP mechanism also influences the short-term response elicited by inotropic agents such as acetylcholine and angiotensin II. A role of NO in long-term cardiac adaptation is illustrated by morphologic evidence (e.g. NOS immuno-localization in phylogenetically distant species) which emphasizes the importance of NO in reshaping the angio-myoarchitecture of the fish heart ventricle (i.e. compensation for regional heterogeneity). Finally, by studying the avascular hearts of teleosts and amphibians that lack vascular endothelium, a relevant role of endocardial endothelium-NO signalling in intracavitary regulation of myocardial performance has been firmly established, thus revealing its early evolutionary role in non-mammalian vertebrates.     

Factorial aerobic scope is independent of temperature and primarily modulated by heart rate in exercising Murray cod (Maccullochella peelii peelii)

Several previous reports, often from studies utilising heavily instrumented animals, have indicated that for teleosts, the increase in cardiac output (Vb) during exercise is mainly the result of an increase in cardiac stroke volume (V(S)) rather than in heart rate (fH). More recently, this contention has been questioned following studies on animals carrying less instrumentation, though the debate continues. In an attempt to shed more light on the situation, we examined the heart rates and oxygen consumption rates (Mo2; normalised to a mass of 1 kg, given as Mo2kg) of six Murray cod (Maccullochella peelii peelii; mean mass+/-SE = 1.81+/-0.14 kg) equipped with implanted fH and body temperature data loggers. Data were determined during exposure to varying temperatures and swimming speeds to encompass the majority of the biological scope of this species. An increase in body temperature (Tb) from 14 degrees C to 29 degrees C resulted in linear increases in Mo2kg (26.67-41.78 micromol min(-1) kg(-1)) and fH (22.3-60.8 beats min(-1)) during routine exercise but a decrease in the oxygen pulse (the amount of oxygen extracted per heartbeat; 1.28-0.74 micromol beat(-1) kg(-1)). During maximum exercise, the factorial increase in Mo2kg was calculated to be 3.7 at all temperatures and was the result of temperature-independent 2.2- and 1.7-fold increases in fH and oxygen pulse, respectively. The constant factorial increases in fH and oxygen pulse suggest that the cardiovascular variables of the Murray cod have temperature-independent maximum gains that contribute to maximal oxygen transport during exercise. At the expense of a larger factorial aerobic scope at an optimal temperature, as has been reported for species of salmon and trout, it is possible that the Murray cod has evolved a lower, but temperature-independent, factorial aerobic scope as an adaptation to the largely fluctuating and unpredictable thermal climate of southeastern Australia.     

Ca2+ protection from the negative inotropic effect of contraction frequency on teleost hearts

Summary Isometric tension development by ventricular strips of 9 species of teleosts, a frog and a turtle was assessed at varying contraction frequencies and Ca_o (external calcium concentration). With teleost hearts an increase in contraction frequency at constant Ca_o was always associated with a decrease in tension development; however, under comparable conditions a positive staircase was exhibited by the frog and turtle heart preparations. The reaction of the teleost heart was thus very different from the well established response of the hearts of higher vertebrates. Elevations in Ca_o always resulted in an increase in tension development such that the positive inotropic effect of Ca_o could compensate for the negative effect of a high contraction frequency. Perfused isolated cod hearts exhibited an increase in cardiac output and pressure development as a result of increases in Ca_o. At 30 contractions min⁻¹ a transition from 1–2 mM Ca_o led to a 68% increase in performance defined as the product of cardiac output times pressure development. The response was in excess of that of ventricular strips. At low Ca_o increases in rate from in situ resting levels to the high end of the physiological range resulted in a decrease in performance. Increases in Ca_o were able to ameliorate the detrimental effect of high imposed contraction frequency. In conclusion, both ventricular strip and perfused heart experiments show that a positive inotropic effect of increased Ca_o can compensate for or even surpass the negative effect of high contraction frequency when both variables are at physiological levels. This finding could have relevance to the maintenance of cardiac performance during/or following intense swimming when both heart rate and plasma calcium may be elevated.     

A quantitative autoradiographic study of 125I atrial natriuretic factor in the heart of a teleost fish (Conger conger).

Quantitative receptor autoradiographic study of 125I-atrial natriuretic peptide factor (ANF) in the heart of a teleost fish Conger conger has shown that a heterogenous distribution of 125I-ANF binding exists in the different cardiac regions. Elevated ANF binding densities (3,790 fmol/mg protein) were encountered in the innermost layer (tunica intima) of the bulbus arteriosus while lower binding levels (293-403 fmol/mg protein) were revealed in atrium and ventricle. In order to determine 125I-ANF binding characteristics (KD, Bmax) in the above cardiac sites, saturation binding assays were carried out. The results show that low 125I-ANF KD values (28.8-52.6 pM) were found in the atrium and in the bulbus arteriosus with respect to the higher KD values (373 pM) of the ventricle. The number of binding sites were respectively 632 and 1,279 fmol/mg protein for the atrium and the ventricle, while a substantially elevated Bmax of 7,235 fmol/mg protein was found for the bulbus arteriosus. These results may furnish some insights concerning ANF receptor binding activity and its putative regulatory role of different cardiac functions.     

Purinergic modulation of cardiac activity in the flounder during hypoxic stress

Summary The interaction of adrenaline and adenosine was examined in cardiac tissue of the flounderPlatichthys flesus.When applied alone both agents increased contractility in both auricular and ventricular myocardial strips. This positive inotropic effect was associated with a small depolarization in the tissues examined by the sucrose gap technique. Simultaneous application of adrenaline and adenosine gave an inhibition of the control responses seen with either agent alone in both auricle and ventricle.Radiocalcium flux studies on ventricular tissue showed that influx was increased by adrenaline or adenosine alone above control values, but when applied together radiocalcium influx was reduced. Radiocalcium efflux from cardiac microsomes was stimulated by challenge with adrenaline or adenosine alone. This stimulation was not seen following simultaneous challenge by both agents.The effect of adrenaline on responses of hypoxic flounder hearts was less than that seen in normoxic hearts. This situation was reversed by pretreatment with the purine receptor blocker caffeine. Caffeine pretreatment also reduced the positive inotropic effect seen in normoxic hearts challenged with adenosine.TLC studies gave strong evidence that hearts perfused with hypoxic salines produced both adenosine and adrenaline.The results are discussed as evidence for a mechanism of heart regulation which the flounder may use as a defence against severe acute hypoxic stress.     

Hemodynamic observations following orthotopic cardiac transplantation: hemodynamic responses to upright exercise at 1 year.

Central hemodynamic responses during upright exercise were studied at 1 year in 40 orthotopic cardiac transplant recipients. Hemodynamic responses were characterized by slow rise in heart rate and blunted peak exercise heart rate response, a significant early increase in stroke index followed by a plateau phase, and a steady increase in ventricular filling pressures and pulmonary artery pressure. In spite of exclusive utilization of the Frank-Starling mechanism to augment cardiac output during early exercise, the pressure responses were comparable to those reported in normal subjects. Our observations also indicate that similarly to normal subjects, the heart rate response plays an important role in the cardiac output achieved at maximum exercise. Although patients with younger donor hearts achieved a more favorable maximum heart rate, the other hemodynamic parameters showed no correlation with the donor heart age. Thus, no hemodynamic disadvantage of older donor hearts could be demonstrated. These data provide further enlightenment regarding the mechanisms of the well-preserved functional capacity noted in these patients.     

Functional morphology of the bulbus arteriosus of rainbow trout (Salmo gairdneri Richardson)

The rate of expansion in volume of the bulbus arteriosus with increase in pressure is measured. From this it is calculated the elastic rebound of the bulbus can account for approximately 25 % of blood flow in the ventral aorta; this proportion decreases as cardiac output increases. The structure of the wall of the bulbus is shown to consist of a compact outer layer with a series of separate longitudinal elements on the inner surface. These elements are connected to the compact layer by numerous radial fibres. This structure equalizes strain in all the structural elements of the wall during large changes in volume. Evidence is discussed which shows that the bulbus arteriosus in teleosts is morphologically and biochemically distinct from the ventral aorta. The bulbus is probably of cardiac origin rather than an expansion of the posterior end of the aorta as generally supposed.     

jumonji gene is essential for the neurulation and cardiac development of mouse embryos with a C3H/He background.

The recessive mutant mouse jumonji (jmj), obtained by a gene trap strategy, shows neural tube defects in approximately half of homozygous embryos with a BALB/cA and 129/Ola mixed background, but no neural tube defects with BALB/cA, C57BL/6J, and DBA/2J backgrounds. Here, we show that neural tube and cardiac defects are observed in all embryos with a C3H/HeJ background. In addition, abnormal groove formation and prominent flexure are observed on the neural plate with full penetrance, suggesting that abnormal groove formation leads to neural tube defects. We found morphogenetic abnormalities in the bulbus cordis (future outflow tract and the right ventricle) of homozygous embryo hearts. Moreover, myocytes in the ventricular trabeculae show hyperplasia with cells filling the ventricles. Together with the observation that the jmj gene is expressed in the neural epithelium of the head neural plate and in myocytes in the bulbus cordis and trabeculae, the results show that the jmj gene plays essential roles in the normal development of the neural plate, morphogenesis of bulbus cordis, and proliferation of trabecular myocytes on a C3H/He background.     

Cardiac natriuretic peptides: a physiological lineage of cardioprotective hormones?

Vertebrate hearts from fish to mammals secrete peptide hormones with profound natriuretic, diuretic, and vasodilatory activity; however, the specific role of these cardiac natriuretic peptides (NPs) in homeostasis is unclear. NPs have been suggested to be involved in salt excretion in saltwater teleosts, whereas they are proposed to be more important in volume regulation in mammals. In this review, we consider an alternative (or perhaps complementary) function of NPs to protect the heart. This hypothesis is based on a number of observations. First, evidence for NPs, or NP-like activity has been found in all vertebrate hearts thus far examined, from osmoconforming saltwater hagfish to euryhaline freshwater and saltwater teleosts to terrestrial mammals. Thus the presence of cardiac NPs appears to be independent of environmental conditions that may variously affect salt and water balance. Second, cardiac stretch is a universal, and one of the most powerful, NP secretagogues. Furthermore, stretch-induced NP release in euryhaline teleosts appears relatively independent of ambient salinity. Third, excessive cardiac stretch that increases end-diastolic volume (EDV) can compromise the mechanical ability of the heart by decreasing actin-myosin interaction (length-tension) or through Laplace effects whereby as EDV increases, the wall tension necessary to maintain a constant pressure must also increase. Excessive cardiac stretch can be produced by factors that decrease cardiac emptying (i.e., increased arterial pressure), or by factors that increase cardiac filling (i.e., increased blood volume, increased venous tone, or decreased venous compliance). Fourth, the major physiological actions of cardiac NPs enhance cardiac emptying and decrease cardiac filling. In fish, NPs promote cardiac emptying by decreasing gill vascular resistance, thereby lowering ventral aortic pressure. In mammals a similar effect is achieved through pulmonary vasodilation. NPs also decrease cardiac filling by decreasing blood volume and increasing venous compliance, the latter producing a rapid fall in central venous pressure. Fifth, the presence of NP clearance receptors in the gill and lung (between the heart and systemic circulation) suggest that these tissues may be exposed to considerably higher NP titers than are systemic tissues. Thus, a decrease in outflow resistance immediately downstream from the heart may be the first response to increased cardiac distension. Because the physiology of cardiac NPs is basically the same in fish and mammals, we propose that the cardioprotective effects of NPs have been well preserved throughout the course of vertebrate evolution. It is also likely that the cardioprotective role of NPs was one of the most primordial homeostatic activities of these peptides in the earliest vertebrates.     

Cloning of a calcitonin gene-related peptide from genomic DNA and its mRNA expression in flounder, Paralichthys olivaceus

A part of genomic DNA including the calcitonin gene-related peptide (CGRP) gene was cloned from flounder by the genome-walking method. The intron/exon boundary was predicted to occur exactly at the same position as in salmon. The 37-amino acid molecule coded by the region from the intron/exon boundary to the stop codon was preceded by a typical Lys-Arg cleavage signal and included a cleavage/amidation site common to the CGRP of other vertebrates. The predicted amino acid sequence of flounder CGRP had 78%, 78%, 78%, 81%, and 73-78% identity to that of salmon, cod, frog, chicken, and mammalian CGRPs, respectively. Among vertebrates, CGRP is more conserved than calcitonin (CT) because the identity of flounder CT to mammalian CTs is 31-50%. Expression analysis indicated that this hormone is synthesized in the brain, heart, intestine, testis, and ovary. Since we have previously shown that the CGRP receptor is expressed in these tissues, it is suggested that CGRP secreted from each tissue functions in a paracrine or autocrine manner.     

Role of creatine phosphokinase in the energy supply of the pumping function of the heart]

The cardiac output of isolated working rat heart and left ventricular pressure were estimated in either almost complete inhibition of creatine kinase by iodoacetamide or predominant fall in adenine nucleotides (AdN) content induced by 2-deoxyglucose treatment. In the former case, a profound cardiac pump failure was observed despite almost normal levels of myocardial AdN and phosphocreatine. Those hearts could not maintain the aortic output at standard load due to lower LV systolic pressure, that was accompanied by increased minimal and maximal diastolic pressures by 5-7 mm Hg as well as by LV diastolic stiffness. As LV systolic pressure in those hearts was unchanged in retrogradely perfused and unloaded hearts it might be suggested that the cardiac pump failure was caused by the decreased LV distensibility. On the contrary, deoxyglucose treatment that resulted in 70% fall in the AdN content was accompanied by only moderate reduction of the cardiac output and insignificant changes in LV diastolic pressure and stiffness. The results suggested that creatine kinase plays a crucial role in the maintenance of normal myofibrillar compliance, which is necessary for cardiac filling and pump function.     

Cardiac performance in Salmo salar with infectious salmon anaemia (ISA): putative role of nitric oxide

Infectious salmon anaemia (ISA) is a viral disease which targets the vascular and endocardial endothelial cells. An in vitro preparation of the spontaneously beating working heart of Salmo salar L. (i.e. able to generate physiological values of output pressure, cardiac output, ventricle work and power) was used to study cardiac performance under basal (i.e. in the absence of stimuli) and loading (i.e. Frank-Starling response) conditions in both control and ISAV-affected fish. In contrast to control fish, the heart preparations of the infected counterparts showed an impairment of the Frank-Starling response, particularly evident in fish infected with a higher virus dose. The Frank-Starling response was progressively impaired with the progression of the viral disease from the time of the virus administration until the 20th day. The potential involvement of nitric oxide (NO) in cardiac dysfunction was investigated by using the authentic nitric oxide synthase (NOS) substrate L-arginine and the NOS inhibitors L-NMMA and L-NIL. In contrast to control fish, infected hearts were particularly sensitive to the inducible nitric oxide synthase (iNOS) inhibitor L-NIL and insensitive to L-arginine. While pretreatment with NOS inhibitors reduced the Frank-Starling response in control hearts, it restored this response in infected counterparts. Taken together, the results indicate that cardiac dysfunction and the NO-transduction-pathway can be mechanistically linked in infected salmon.     

The right ventricular working heart preparation

An isolated working rat heart preparation was modified to study right ventricular (RV) performance. All hearts were perfused with a Krebs-Henseleit bicarbonate buffer via a Langendorff column at 90 mm Hg. Right atrial filling (preload) was varied by raising a buffer reservoir from 5 cm below to 10 cm above the right atrium while pulmonary artery outflow resistance remained fixed. RV systolic pressure and the maximum rise and decrease in pressure development (+/- dP/dt) were measured via a catheter in the RV. Cardiac output was collected with a catheter placed in the pulmonary artery. One group of hearts, monitored at a fixed preload (0 cm H2O) for 2 hr, and another group of hearts, in which two ventricular function curves were performed, demonstrated the stability and reproducibility of the preparation. Additionally, the ability of this preparation to measure changes in inotropy was studied. A negative inotropic effect was measured after verapamil (5 X 10(-8) M) treatment. Positive dP/dt showed the greatest depression (30%) and was significantly lower at every preload. A positive inotropic effect was demonstrated by reducing the buffer Ca2+ concentration to 1.9 mM for the first work curve followed by an addition of Ca2+ (2.8 mM final concentration) or ouabain (5 X 10(-5) M) for the second work curve. Again, the greatest effect was found in the dP/dt measurements (elevated by 20 and 30%, respectively). Thus, this preparation manifests qualities similar to those used in studying the left ventricle and allows investigation of various cardiac diseases which may affect RV pump function.