Control of heart rate during thermoregulation in the heliothermic lizard Pogona barbata: importance of cholinergic and adrenergic mechanisms

During thermo regulation in the bearded dragon Pogona barbata, heart rate when heating is significantly faster than when cooling at any given body temperature (heart rate hysteresis), resulting in faster rates of heating than cooling. However, the mechanisms that control heart rate during heating and cooling are unknown. The aim of this study was to test the hypothesis that changes in cholinergic and adrenergic tone on the heart are responsible for the heart rate hysteresis during heating and cooling in P. barbata. Heating and cooling trials were conducted before and after the administration of atropine, a muscarinic antagonist, and sotalol, a beta-adrenergic antagonist. Cholinergic and beta-adrenergic blockade did not abolish the heart rate hysteresis, as the heart rate during heating was significantly faster than during cooling in all cases. Adrenergic tone was extremely high (92.3%) at the commencement of heating, and decreased to 30.7% at the end of the cooling period. Moreover, in four lizards there was an instantaneous drop in heart rate (up to 15 beats min(-1)) as the heat source was switched off, and this drop in heart rate coincided with either a drop in beta-adrenergic tone or an increase in cholinergic tone. Rates of heating were significantly faster during the cholinergic blockade, and least with a combined cholinergic and beta-adrenergic blockade. The results showed that cholinergic and beta-adrenergic systems are not the only control mechanisms acting on the heart during heating and cooling, but they do have a significant effect on heart rate and on rates of heating and cooling.

Temperature and the respiratory properties of whole blood in two reptiles, Pogona barbata and Emydura signata

We investigated the capacity of two reptiles, an agamid lizard Pogona barbata and a chelid turtle Emydura signata, to compensate for the effects of temperature by making changes in their whole blood respiratory properties. This was accomplished by measuring the P-50 (at 10, 20 and 30 degrees C), hematocrit (Hct), haemoglobin concentration ([Hb]) and mean cell haemoglobin concentration (MCHC) in field acclimatised and laboratory acclimated individuals. The acute effect of temperature on P50 in P barbata, expressed as heat of oxygenation (Delta H), ranged from -16.8 +/- 1.84 to -28.5 +/- 2.73 kJ/mole. P-50 of field acclimatised P barbata increased significantly from early spring to summer at the test temperatures of 20 degrees C (43.1 +/- 1.2 to 48.8 +/- 2.1 mmHg) and 30 degrees C (54.7 +/- 1.2 to 65.2 +/- 2.3 mmHg), but showed no acclimation under laboratory conditions. For E. signata, Delta H ranged from -31.1 +/- 6.32 to -48.2 +/- 3.59 kJ/mole. Field acclimatisation and laboratory acclimation of P-50 did not occur. However, in E. signata, there was a significant increase in [Hb] and MCHC from early spring to summer in turtles collected from the wild (1.0 +/- 0.1 to 1.7 +/- 0.2 mmol/L and 4.0 +/- 0.3 to 6.7 +/- 0.7 mmol/L, respectively). (C) 2005 Published by Elsevier Inc.

The effect of heat transfer mode on heart rate responses and hysteresis during heating and cooling in the estuarine crocodile Crocodylus porosus

The effect of heating and cooling on heart rate in the estuarine crocodile Crocodylus porosus was studied in response to different heat transfer mechanisms and heat loads. Three heating treatments were investigated. C. porosus were: (1) exposed to a radiant heat source under dry conditions; (2) heated via radiant energy while half-submerged in flowing water at 23degreesC and (3) heated via convective transfer by increasing water temperature from 23degreesC to 35degreesC. Cooling was achieved in all treatments by removing the heat source and with C. porosus half-submerged in flowing water at 23degreesC. In all treatments, the heart rate of C. porosus increased markedly in response to heating and decreased rapidly with the removal of the heat source. Heart rate during heating was significantly faster than during cooling at any given body temperature, i.e. there was a significant heart rate hysteresis. There were two identifiable responses to heating and cooling. During the initial stages of applying or removing the heat source, there was a dramatic increase or decrease in heart rate ('rapid response'), respectively, indicating a possible cardiac reflex. This rapid change in heart rate with only a small change or no change in body temperature (

Integration of autonomic and local mechanisms in regulating cardiovascular responses to heating and cooling in a reptile (Crocodylus porosus)

Reptiles change heart rate and blood flow patterns in response to heating and cooling, thereby decreasing the behavioural cost of thermoregulation. We tested the hypothesis that locally produced vasoactive substances, nitric oxide and prostaglandins, mediate the cardiovascular response of reptiles to heat. Heart rate and blood pressure were measured in eight crocodiles (Crocodylus porosus) during heating and cooling and while sequentially inhibiting nitric-oxide synthase and cyclooxygenase enzymes. Heart rate and blood pressure were significantly higher during heating than during cooling in all treatments. Power spectral density of heart rate and blood pressure increased significantly during heating and cooling compared to the preceding period of thermal equilibrium. Spectral density of heart rate in the high frequency band (0.19-0.70 Hz) was significantly greater during cooling in the saline treatment compared to when nitric-oxide synthase and cyclooxygenase enzymes were inhibited. Cross spectral analysis showed that changes in blood pressure preceded heart rate changes at low frequencies (

Physiological thermoregulation in a crustacean? Heart rate hysteresis in the freshwater crayfish Cherax destructor

Differential heart rates during heating and cooling (heart rate hysteresis) are an important thermoregulatory mechanism in ectothermic reptiles. We speculate that heart rate hysteresis has evolved alongside vascularisation, and to determine whether this phenomenon occurs in a lineage with vascularised circulatory systems that is phylogenetically distant from reptiles, we measured the response of heart rate to convective heat transfer in the Australian freshwater crayfish, Cherax destructor. Heart rate during convective heating (from 20 to 30 degreesC) was significantly faster than during cooling for any given body temperature. Heart rate declined rapidly immediately following the removal of the heat source, despite only negligible losses in body temperature. This heart rate 'hysteresis' is similar to the pattern reported in many reptiles and, by varying peripheral blood flow, it is presumed to confer thermoregulatory benefits particularly given the thermal sensitivity of many physiological rate functions in crustaceans. (C) 2004 Published by Elsevier Inc.

Diving behaviour of a reptile (Crocodylus johnstoni) in the wild: Interactions with heart rate and body temperature

The differences in physical properties of air and water pose unique behavioural and physiological demands on semiaquatic animals. The aim of this study was to describe the diving behaviour of the freshwater crocodile Crocodylus johnstoni in the wild and to assess the relationships between diving, body temperature, and heart rate. Time-depth recorders, temperature-sensitive radio transmitters, and heart rate transmitters were deployed on each of six C. johnstoni (4.0-26.5 kg), and data were obtained from five animals. Crocodiles showed the greatest diving activity in the morning (0600-1200 hours) and were least active at night, remaining at the water surface. Surprisingly, activity pattern was asynchronous with thermoregulation, and activity was correlated to light rather than to body temperature. Nonetheless, crocodiles thermoregulated and showed a typical heart rate hysteresis pattern (heart rate during heating greater than heart rate during cooling) in response to heating and cooling. Additionally, dive length decreased with increasing body temperature. Maximum diving length was 119.6 min, but the greatest proportion of diving time was spent on relatively short (

Physiological mechanisms of thermoregulation in reptiles: a review

The thermal dependence of biochemical reaction rates means that many animals regulate their body temperature so that fluctuations in body temperature are small compared to environmental temperature fluctuations. Thermoregulation is a complex process that involves sensing of the environment, and subsequent processing of the environmental information. We suggest that the physiological mechanisms that facilitate thermoregulation transcend phylogenetic boundaries. Reptiles are primarily used as model organisms for ecological and evolutionary research and, unlike in mammals, the physiological basis of many aspects in thermoregulation remains obscure. Here, we review recent research on regulation of body temperature, thermoreception, body temperature set-points, and cardiovascular control of heating and cooling in reptiles. The aim of this review is to place physiological thermoregulation of reptiles in a wider phylogenetic context. Future research on reptilian thermoregulation should focus on the pathways that connect peripheral sensing to central processing which will ultimately lead to the thermoregulatory response.

Prostaglandins are important in thermoregulation of a reptile (Pogona vitticeps)

The effectiveness of behavioural thermoregulation in reptiles is amplified by cardiovascular responses, particularly by differential rates of heart beat in response to heating and cooling (heart-rate hysteresis). Heart-rate hysteresis is ecologically important in most lineages of ectothermic reptile' and we demonstrate that heart-rate hysteresis in the lizard Pogona vitticeps is mediated by prostaglandins. In a control treatment (administration of saline), heart rates during heating were significantly faster than during cooling at any given body temperature. When cyclooxygenase 1 and 2 enzymes were inhibited, heart rates during heating were not significantly different from those during cooling. Administration of agonists showed that thromboxane B-2 did not have a significant effect on heart rate, but prostacyclin and prostaglandin F-2alpha caused a significant increase (3.5 and 13.6 beats min(-1), respectively) in heart rate compared with control treatments. We speculate that heart-rate hysteresis evolved as a thermoregulatory mechanism that may ultimately be controlled by neurally induced stimulation of nitric oxide production, or maybe via photolytically induced production of vitamin D.

Level and precision of behavioural thermoregulation in the bearded dragon, Pogona vitticeps : effects of hypoxia and environmental thermal quality /

Most metabolic functions are optimized within a narrow range of body temperatures, which is why thermoregulation is of great importance for the survival and overall fitness of an animal. It has been proposed that lizards will thermoregulate less precisely in low thermal quality environments, where the costs associated with thermoregulation are high; in the case of lizards, whose thermoregulation is mainly behavioural, the primary costs ofthermoregulation are those derived from locomotion. Decreasing thermoregulatory precision in costly situations is a strategy that enhances fitness by allowing lizards to be more flexible to changing environmental conditions. It allows animals to maximize the benefits of maintaining a relatively high body temperature while minimizing energy expenditure. In situations where oxygen concentration is low, the costs of thermoregulation are relatively high (i.e. in relation to the amount of oxygen available for metabolic functions). As a result, it is likely that exposures to hypoxic conditions induce a decrease in the precision of thermoregulation. This study evaluated the effects of hypoxia and low environmental thermal quality, two energetically costly conditions, on the precision and level of thermoregulation in the bearded dragon, Pogona vitticeps, in an electronic temperature-choice shuttle box. Four levels of hypoxia (1O, 7, 5 and 4% 02) were tested. Environmental thermal quality was manipulated by varying the rate of temperature change (oTa) in an electronic temperature-choice shuttle box. Higher oT a's translate into more thermally challenging environments, since under these conditions the animals are forced to move a greater number of times (and hence invest more energy in locomotion) to maintain similar temperatures than at lower oTa's. In addition, lizards were tested in an "extreme temperatures" treatment during which air temperatures of the hot and cold compartments of the shuttle box were maintained at a constant 50 and 15°C respectively. This was considered the most thermally challenging environment. The selected ambient (T a) and internal body temperatures (Tb) of bearded dragons, as well as the thermoregulatory precision (measured by the central 68% ofthe Ta and T b distribution) were evaluated. The thermoregulatory response was similar to both conditions. A significant increase in the size of the Tb range, reflecting a decrease in thermoregulatory precision, and a drop in preferred body temperature of ~2 °C, were observed at both 4% oxygen and at the environment of lowest thermal quality. The present study suggests that in energetically costly situations, such as the ones tested in this study, the bearded dragon reduces energy expenditure by decreasing preferred body temperature and minimizing locomotion, at the expense of precise behavioural thermoregulation. The close similarity of the behavioural thermoregulatory response to two very different stimuli suggests a possible common mechanism and neuronal pathway to the thermoregulatory response.

Thermoregulatory consequences of salt loading in the lizard Pogona vitticeps

Previous research has demonstrated that dehydration increases the threshold temperature for panting and decreases the thermal preference of lizards. Conversely, it is unknown whether thermoregulatory responses such as shuttling and gaping are similarly influenced. Shuttling, as an active behavioural response, is considered one of the most effective thermoregulatory behaviours, whereas gaping has been proposed to be involved in preventing brain over-heating in lizards. In this study we examined the effect of salt loading, a proxy for increased plasma osmolality, on shuttling and gaping in Pogona vitticeps. Then, we determined the upper and lower escape ambient temperatures (UETa and LETa), the percentage of time spent gaping, the metabolic rate ((V) over dot(O2)), the evaporative water loss (EWL) during gaping and non-gaping intervals and the evaporative effectiveness (EWL/(V) over dot(O2)) of gaping. All experiments were performed under isotonic (154 mmol l(-1)) and hypertonic saline injections (625, 1250 or 2500 mmol l(-1)). Only the highest concentration of hypertonic saline altered the UETa and LETa, but this effect appeared to be the result of diminishing the animal's propensity to move, instead of any direct reduction in thermoregulatory set-points. Nevertheless, the percentage of time spent gaping was proportionally reduced according to the saline concentration; (V) over dot(O2) was also decreased after salt loading. Thermographic images revealed lower head than body surface temperatures during gaping; however this difference was inhibited after salt loading. Our data suggest that EWL/(V) over dot(O2) is raised during gaping, possibly contributing to an increase in heat transfer away from the lizard, and playing a role in head or brain cooling.

Influencia de las poliaminas en la deposición del carbonato cálcico en el alga Cymopolia barbata (Chlorophyta, Dasycladales).

Papel de las poliaminas en la precipitación del carbonato en algas

Habitat shifts: an experimental study on disappearing canopies of Cystoseira barbata (Stackhouse) C. Agard and the habitat replacing them

Cambiamenti di habitat in ambienti marini: uno studio sperimentale sulla perdita di foreste a Cystoseira barbata (Stackhouse) C. Agardh e sui popolamenti che le sostituiscono La presente tesi affronta il tema scientifico generale di come prevedere e mitigare la perdita di habitat marini naturali causata dalle attività umane. La tesi si è focalizzata sugli habitat subtidali a “canopy” formati da macroalghe brune a tallo eretto dell’ordine Fucales, che per morfologia, ruolo ed importanza ecologica possono essere paragonate alle “foreste” in ambienti terrestri temperati. Questi sistemi sono tra i più produttivi in ambienti marini, e sono coinvolti in importanti processi ecologici, offrendo cibo, protezione, riparo ed ancoraggio a diverse altre specie animali e vegetali, modificando i gradienti naturali di luce, sedimentazione e idrodinamismo, e partecipando al ciclo dei nutrienti. Sulle coste temperate di tutto il mondo, le foreste di macroalghe a canopy sono in forte regressione su scala locale, regionale e globale. Questo fenomeno, che sta accelerando a un ritmo sempre più allarmante, sta sollevando interesse e preoccupazione. Infatti, data la loro importanza, la perdita di questi habitat può avere importanti conseguenze ecologiche ed economiche, tra cui anche il possibile declino della pesca che è stato osservato in alcune aree in seguito alla conseguente riduzione della produttività complessiva dei sistemi marini costieri. Nel Mar Mediterraneo questi tipi di habitat sono originati prevalentemente da alghe appartenenti al genere Cystoseira, che sono segnalate in forte regressione in molte regioni. Gli habitat a Cystoseira che ancora persistono continuano ad essere minacciati da una sineregia di impatti antropici, ed i benefici complessivi delle misure di protezione fin ora attuate sono relativamente scarsi. Scopo della presente tesi era quello di documentare la perdita di habitat a Cystoseira (prevalentemente Cystoseira barbata (Stackhouse) C. Agardh) lungo le coste del Monte Conero (Mar Adriatico centrale, Italia), e chiarire alcuni dei possibili meccanismi alla base di tale perdita. Studi precedentemente condotti nell’area di studio avevano evidenziato importanti cambiamenti nella composizione floristica e della distribuzione di habitat a Cystoseira in quest’area, e avevano suggerito che la scarsa capacità di recupero di questi sistemi potesse essere regolata da interazioni tra Cystoseira e le nuove specie dominanti sui substrati lasciati liberi dalla perdita di Cystoseira. Attraverso ripetute mappature dell’habitat condotte a partire da Luglio 2008 fino a Giugno 2010, ho documentato la perdita progressiva delle poche, e sempre più frammentate, patch di habitat originate da questa specie in due siti chiamati La Vela e Due Sorelle. Attraverso successivi esperimenti, ho poi evidenziato le interazioni ecologiche tra le specie dominanti coinvolte in questi cambiamenti di habitat, al fine di identificare possibili meccanismi di feedback che possano facilitare la persistenza di ciascun habitat o, viceversa, l’insediamento di habitat alternativi. La mappatura dell’habitat ha mostrato un chiaro declino della copertura, della densità e della dimensione degli habitat a Cystoseira (rappresentati soprattutto dalla specie C. barbata e occasionalmente C. compressa che però non è stata inclusa nei successivi esperimenti, d’ora in avanti per semplicità verrà utilizzato unicamente il termine Cystoseira per indicare questo habitat) durante il periodo di studio. Nel sito Due Sorelle le canopy a Cystoseira sono virtualmente scomparse, mentre a La Vela sono rimaste poche, sporadiche ed isolate chiazze di Cystoseira. Questi habitat a canopy sono stati sostituiti da nuovi habitat più semplici, tra cui soprattutto letti di mitili, feltri algali e stand monospecifici di Gracilaira spp.. La mappatura dell’habitat ha inoltre sottolineato una diminuzione del potenziale di recupero del sistema con un chiaro declino del reclutamento di Cystoseira durante tutto il periodo di studio. Successivamente ho testato se: 1) una volta perse, il recupero di Cystoseira (reclutamento) possa essere influenzato dalle interazioni con le nuove specie dominanti, quali mitili e feltri algali; 2) il reclutamento di mitili direttamente sulle fronde di Cystoseira (sia talli allo stadio adulto che giovanili) possa influenzare la sopravvivenza e la crescita della macroalga; 3) la sopravvivenza e la crescita delle nuove specie dominanti, in particolare mitili, possa essere rallentata dalla presenza di canopy di Cystoseira. I risultati dimostrano che le nuove specie dominanti insediatesi (feltri algali e mitili), possono inibire il reclutamento di Cystoseira, accelerandone il conseguente declino. L’effetto diretto dei mitili sulle fronde non è risultato particolarmente significativo né per la sopravvivenza di Cystoseira che finora non è risultata preclusa in nessun stadio di sviluppo, né per la crescita, che nel caso di individui adulti è risultata leggermente, ma non significativamente, più elevata per le fronde pulite dai mitili, mentre è stato osservato il contrario per i giovanili. La presenza di canopy a Cystoseira, anche se di piccole dimensioni, ha limitato la sopravvivenza di mitili. Questi risultati complessivamente suggeriscono che una foresta di macroalghe in buone condizioni può avere un meccanismo di autoregolazione in grado di facilitare la propria persistenza. Quando però il sistema inizia a degradarsi e a frammentarsi progressivamente, i cambiamenti delle condizioni biotiche determinati dall’aumento di nuove specie dominanti contribuiscono alla mancanza di capacità di recupero del sistema. Pertanto le strategie per una gestione sostenibile di questi sistemi dovrebbero focalizzarsi sui primi segnali di cambiamenti in questo habitat e sui possibili fattori che ne mantengono la resilienza.

Campanula abietina - Dracocephalum ruyschiana - Pentstemon - Campanula persicifolia - Campanula barbata - Campanula rhomboidalis

AR

Recovery plan for two insects (Polyphylla barbata and Trimerotropis infantilis) and four plants (Chorizanthe pungens var. hartwegiana, Chorizanthe robusta var. hartwegii, Erysimum teretifolium, and Polygonum hickmanii) from the Santa Cruz mountains : technical and agency draft /

"August 1997."