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Circadian organization in ruin lizards: phase response curve for melatonin changes with season.

Single biweekly injections of exogenous melatonin were given to ruin lizards ( Podarcis sicula) whose circadian locomotor rhythms were recorded under constant temperature (29 degrees C) and darkness. To generate a phase-response curve the direction and amount of phase shift were plotted versus the time at which the injection was given. The results showed, for the second time in the animal kingdom, the existence of a phase-response curve for melatonin. The area under the delay part of the phase-response curve is much greater than the area under the advance part, and there is no dead zone. The phase-response curve for melatonin was generated in summer, but the repetition of experiments in autumn-winter did not generate any phase-response curve for melatonin in these seasons. In autumn-winter no one phase of the ruin lizards' circadian cycle was shifted in response to melatonin injections. This confirms several previous results in ruin lizards, showing that melatonin is centrally involved in determining circadian organization in summer and only marginally involved in autumn-winter.

Animals↗

Whole-organism studies of adhesion in pad-bearing lizards: creative evolutionary solutions to functional problems.

Understanding the evolution of complex functional traits is a challenge for evolutionary physiology. Here we investigate the evolution of subdigital toepads in lizards, which have arisen independently at least three times, although with subtle anatomical differences. Some designs (anole, gecko) appear functionally equivalent, whereas other designs (skink) are inferior. The functional equivalence of geckos and anoles highlights the creative aspect of the evolutionary process in that these two groups have arrived at the same functional endpoint along very different trajectories. However, this functional equivalence does not result in equivalence for performance at whole-organism tasks (e.g., running uphill), as the evolution of behavior (e.g., toe-furling) has enabled geckos to be superior climbers than anoles. We also show that adaptive increases in the toepad size within a closely related lizard genus (Anolis) has resulted in concomitant evolution of enhanced clinging ability and increased perch heights. A third insight is that pad-bearing geckos are capable of carrying tremendous loads (up to 250% of body weight) up smooth surfaces, and that the toepad itself does not appear limiting. This comparative and whole-organism approach to lizard toepads underscores how organisms can evolve multiple solutions to evolutionary problems.

Adaptation, Physiological↗

The effect of temperature on digestive and assimilation efficiency, gut passage time and appetite in an ambush foraging lizard, Cordylus melanotus melanotus.

In ectotherms, an increase in body temperature increases metabolic rate and may increase rates of digestive processes. We measured the thermal dependence of the apparent digestive and apparent assimilation efficiencies (ADE and AAE), gut passage time (GP) and appetite in Cordylus melanotus melanotus, a medium sized Crag Lizard, which is endemic to South Africa. Trials were conducted at 20, 22, 25, 30, 32 and 35 degrees C under controlled conditions. Trials lasted 14 days, during which, lizards were fed ca. 1 g mealworms per day. Glass beads were used as markers to determine GP at the beginning and end of trials. Faeces and urates were collected daily and oven dried at 50 degrees C. The energy content of egested matter was then measured using bomb calorimetry. ADE and AAE were not affected by temperature for either males or females. The mean+/-SE ADE and AAE were 94.4+/-0.3% and 87.2+/-0.6%, respectively. GP was not significantly different between males and females at any temperature, but decreased significantly with increasing temperature. Appetite was significantly different between the different temperatures measured. The decrease of gut passage time with increasing temperature was expected, since the digestive and assimilation efficiencies are similar over the range of temperatures tested. Lizards are thus assimilating a similar proportion of ingested energy, but at faster rates at higher temperatures. The results indicate that the digestive physiology of this species results in maximum energy gain per meal in environments where food is scarce.

Animals↗

Prevalence and intensity of haemogregarine blood parasites and their mite vectors in the common wall lizard, Podarcis muralis.

We describe the general field population biology of haemogregarines and mites in the wall lizard, Podarcis muralis, examining the factors that determine the prevalence and intensity of infection. The intensity of infection by haemogregarines in females was slightly lower in summer than in spring, whereas males maintained similar levels of intensity among all the seasons, probably due to immunosuppressive effects of testosterone. This is also supported because the prevalence and the infection intensity by mites were higher in males than in females. Parasite load was higher in lizards with better body condition, which could be reflecting the mortality of infected lizards with poor condition.

Animals↗

Genetic variation in parthenogenetic Caucasian rock lizards of the genus Lacerta (L. dahli, L. armeniaca, L. unisexualis) analyzed by DNA fingerprinting.

Multilocus DNA fingerprinting has been used to study the variability of some mini- and microsatellite sequences in parthenogenetic species of Caucasian rock lizards of the genus Lacerta (L. dahli, L. armeniaca and L. unisexualis). We demonstrate that these clonally reproducing lizards possess species-specific DNA fingerprints with a low degree of intra- and interpopulation variation. Mean indices of similarity obtained using M13 DNA, (GACA)4 and (TCC)50 as probes were 0.962 and 0.966 in L. dahli and L. armeniaca, respectively. The mean index of similarity obtained using M 13 and GATA probes in L. unisexualis was estimated to be 0.95. However, despite the high degree of band-sharing, variable DNA fragments were revealed in all populations with the microsatellite probes. An particularly high level of variability was observed for (TCC)n microsatellites in populations of L. unisexualis. In fact TCC-derived DNA fingerprints were close to being individual-specific, with a mean index of similarity of 0.824. Fingerprint analysis of parthenogenetic families of L. armeniaca showed that all maternal fragments were inherited together by the progeny, and no differences in fingerprint patterns were observed. On the other hand, while identical DNA fingerprints were obtained from L. unisexualis families with M13 and (GATA)4 probes, use of the (TCC)50 probe revealed remarkable intrafamily variation in this species. It is assumed that the genetic heterogeneity observed in parthenogenetic populations may be explained, at least in part, by the existence of genetically unstable microsatellite loci. Our data serve to illustrate processes of spontaneous mutagenesis and the initial stages of clonal differentiation in natural populations of the lizard species studied.

Animals↗

Reliable signaling by chemical cues of male traits and health state in male lizards, Lacerta monticola.

In spite of the importance of chemoreception in social organization and sexual selection of lizards, there is a lack of general knowledge on how the characteristics of chemical signals mediate these behaviors. Moreover, it is unknown which are the mechanisms that might confer honesty to the information provided by chemical signals. We analyzed here whether characteristics of the lipophilic fraction of femoral gland secretions of Lacerta monticola male lizards can be related to the morphology, physical condition, and health state of the sender. Our results indicated that some male traits, such as body size, number of blue spots, and number of femoral pores and their level of fluctuating asymmetry, were related to variability in the relative proportions of some lipophilic chemical compounds found in secretions. Thus, conspecifics could obtain reliable information on the producer of a scent mark based on chemicals alone, and this might be the basis of female choice observed in this lizard. Moreover, only males with a greater T-cell immune response had higher proportions of two steroids (ergosterol and dehydrocholesterol) in their femoral secretions, which might suggest that the signal is honest and costly to produce. We suggest that only high-quality males could divert these compounds from metabolism to secretions in order to produce an exaggerated and honest "chemical ornament."

Animals↗

Individual, sexual, seasonal, and temporal variation in the amount of sagebrush lizard scent marks.

Although many animals deposit scent marks, previous studies have focused almost entirely on rodents or on the chemical structure of the signal. Here, we study the quantity and temporal pattern of chemical deposition by the territorial sagebrush lizard Sceloporus graciosus, measuring both femoral pore and fecal deposits. Specifically, we tested whether variation in deposition is a good cue of individual and sexual identity and/or whether it is more closely associated with body size and reproductive state, indicators of physiological condition. The results support the latter hypothesis. We found that although the amount of fluid deposited on a single perch (rarely quantified in mammals) carries little information on individual or sexual identity, it reflects the physiological condition and reproductive state of individual lizards and is replenished on a roughly weekly cycle, potentially providing additional information on the producer's activity level. The amount of deposition may thus provide important information to chemical receivers making mate choice and territorial defense decisions. The results further suggest that seasonal increases in gland production allow lizards to mark more sites rather than to influence the quality of the signal on a single perch.

Animals↗

Chemically mediated species recognition in closely related Podarcis wall lizards.

In many animals, chemical signals play an important role in species recognition and may contribute to reproductive isolation and speciation. The Iberian lizards of the genus Podarcis, with up to nine currently recognized lineages that are often sympatric, are highly chemosensory and provide an excellent model for the study of chemically mediated species recognition in closely related taxa. In this study, we tested the ability of male and female lizards of two sister species with widely overlapping distribution ranges (Podarcis bocagei and P. hispanica type 1) to discriminate between conspecific and heterospecific mates by using only substrate-borne chemical cues. We scored the number of tongue flicks directed at the paper substrate by each individual in a terrarium previously occupied by a conspecific or a heterospecific lizard of the opposite sex. Results show that males of P. bocagei and P. hispanica type 1 are capable of discriminating chemically between conspecifics and heterospecifics of the opposite sex, but females are not. These results suggest that differences in female, but not male, chemical cues may underlie species recognition and contribute to reproductive isolation in these species. The apparent inability of females to discriminate conspecific from heterospecific males, which is not because of reduced baseline exploration rates, is discussed in the context of sexual selection theory and species discrimination.

Animal Communication↗

Alterations of cell membrane composition during prolonged fasting in lizards.

Male lizards fasted for 35 days showed significant alterations in the lipid composition of membranes isolated from a variety of organs and tissues. Unesterified cholesterol was significantly increased in membranes from kidney, skeletal muscle, small intestine, testis and heart but not in those from lung, brain and liver; cholesterol/phospholipid molar ratio was also significantly increased in membranes from all organs except brain and liver. Membrane total phospholipids were reduced for all tissues except brain and liver, whilst significant reductions in the molar ratio of membrane phosphatidylcholine to sphingomyelin occurred in kidney, muscle, testis, intestine and liver. Similar alterations in lipid composition were found in plasma and significant correlations between plasma and membrane cholesterol/phospholipid and phosphatidylcholine/sphingomyelin molar ratios were found for those organs in which significant lipid changes occurred during fasting. Lizards fasted for 35 days and then refed a natural diet, such that they started to regain weight, had normal membrane lipid compositions. These observations suggest that during prolonged fasting certain tissues of the lizard are unable to maintain a normal membrane lipid composition and that abnormal plasma lipoproteins may be involved in this process.

Animals↗

Localization of neurons containing pro-opiomelanocortin-related peptides in the hypothalamus and midbrain of the lizard, Anolis carolinensis: evidence for region-specific processing of beta-endorphin.

Immunohistochemical analyses of the lizard-brain, following colchicine pretreatment, revealed two populations of POMC-producing cell bodies located in medial-basal hypothalamus and the mesencephalic tegmentum. Analyses of extracts of lizard brain regions by radioimmunoassay and gel filtration chromatography indicate that beta-endorphin-sized and alpha-MSH-sized peptides are the major POMC-related end products. Evidence is presented for region-specific processing of beta-endorphin in the lizard brain.

Animals↗

Serotonin innervation of the cerebral cortex in lizards.

The serotonin (5-HT) innervation of the cerebral cortex in two species of lizards has been studied. Results show no differences between both species. Most of the cerebral cortex of these lizards is innervated by serotoninergic fibers, which are fine and varicose. Their density varies greatly from one cortical region to another: the areas with higher density of serotoninergic fibers and terminals are parts of the medial and dorsal cortices. There is a laminar pattern of distribution of serotoninergic fibers. In the medial cortex, 5-HT fibers are found preferentially in both plexiform layers just above and below the cellular layer. In the dorsomedial cortex, there is an immunoreactive plexus in the outermost third of the superficial plexiform layer and another in the depth of the layer, whereas 5-HT fibers are distributed evenly in the deep plexiform layer of this cortex. In the pars medialis of the dorsal cortex, serotoninergic fibers are abundant in all layers, whereas in the pars lateralis, fibers are found predominantly in the external third of the superficial plexiform layer. The lateral cortex is almost devoid of immunoreactive fibers. These results show a different organization of the cortical serotonin innervation between lizards and turtles.

Animals↗

Parvalbumin-immunoreactive neurons in the cerebral cortex of the lizard Podarcis hispanica.

An antibody against the calcium binding protein parvalbumin selectively labels a set of neurons in the cerebral cortex of lizards. Golgi-like immunostained bipolar, multipolar and pyramid-like neurons appear mainly located in the inner plexiform layers. Parvalbumin-immunoreactive (PARV-IR) puncta are concentrated in the cell layer of the dorsal and dorsomedial cortices showing a basket-like distribution. The morphology and distribution of PARV-IR neurons and puncta overlap GABA-immunostaining in the cerebral cortex of lizards. Thus, it is likely that PARV-IR neurons are a subset of the cortical GABAergic neurons of lizards.

Animals↗

Changes in morphometric parameters and the characteristics of collagen following thyroxine and thiourea treatments in young male garden lizards.

Administration of both thyroxine (1 microgram/g body wt) and thiourea (0.5 mg/10 g body wt) to young male garden lizards for a period of 2 weeks led to changes in morphometric/biochemical parameters. Percentages of increase in body weight and snout-to-vent length of thyroxine-treated lizards during this period were significantly lower than the percentages of increase observed in control lizards. Thiourea-induced inhibition of increase in body weight was marginal but not significant statistically. The total collagen content of tendons decreased upon thyroxine treatment and increased following thiourea administration. Thyroxine treatment led to an increase in salt solubility of bone and skin collagens and thiourea showed a reverse effect. On the other hand, the acid solubility of tendon collagen increased after thyroxine administration and decreased following thiourea treatment.

Aging↗

Effects of TRH on hormone release from pituitaries of the lizard, Anolis carolinensis.

The ability of thyrotropin-releasing hormone (TRH) to stimulate thyrotropin (TSH) from pituitaries of adult male lizards, Anolis carolinensis, was tested in vivo and in vitro. TSH output by pituitaries in vitro was determined by bioassay of the incubation medium using in vitro T4 output by thyroids from the same lizards. Pituitaries incubated without TRH had no detectable TSH secretion during two consecutive (2 or 3 hr) periods of incubation. Incubation in 10 or 100 ng/ml TRH for 2 or 3 hr significantly augmented release of TSH bioactivity in a dose-dependent manner. Pituitaries taken from goitrogen (100 micrograms methimazole/day for 10 days)-treated lizards had elevated basal TSH secretion but did not respond to TRH. TRH injection in vivo (5 micrograms/hr for 10 hr) appeared to stimulate acute release of MSH activity as judged by darkening of skin color after each injection, and plasma T4 was significantly elevated at the end of treatment. These results provide additional evidence that the reptilian thyrotrope has functional TRH receptors and the TSH-stimulating activity of the tripeptide along with its effects on other pituitary cells was present at an early stage of reptilian evolution.

Animals↗

Evidence for the role of thyroxine as a hormone in the physiology of a lizard.

In vivo effects of thyroxine (T4) and triiodothyronine (T3) were studied in male lizards. T4 or T3 (0.5 to 2 nmol) was administered per day over 10 days to surgically thyroidectomized male lizards (35 +/- 2.0 g) and the responses were measured in terms of scale shedding, whole body oxygen consumption (OC), and testicular weight. T3 was more effective in stimulating OC as compared to T4. T4 accelerated scale shedding to a greater extent as compared to T3. T4 was more effective than T3 in restoring the decline in the gonadal weight of thyroidectomized animals. Effect of inhibition of peripheral conversion of T4----T3 by iopanoic acid (IOP) was studied on the above response parameters. IOP at both dose levels inhibited extrathyroidal conversion of T4----T3. The T4-stimulated increase in OC of IOP-treated animals was suppressed, clearly indicating that the T4 effect could be attributed to its conversion to T3. But in these same animals, IOP failed to inhibit T4-stimulated scale shedding and gonadal weight. As the proportion of T4 converted to T3 decreased as a result of IOP treatment, the effectiveness of T4 increased in terms of scale shedding and restoration of gonadal weight. From these studies it appears that all effects of T4 in lizards need not necessarily be mediated via conversion to T3.

Animals↗

Lack of effect of TRH on alpha-MSH release from the neurointermediate lobe of the lizard Lacerta vivipara.

Thyrotropin-releasing hormone (TRH) is a potent stimulator of melanotropin (alpha-MSH) release from pituitary melanotrophs in pig, frog, and fish. Concurrently, it has recently been shown that injection of TRH induces skin darkening in the lizard Anolis carolinensis (Licht and Denver, 1988). In the present study, we have thus investigated in vitro the possible effect of TRH on alpha-MSH release from the lizard (Lacerta vivipara) neurointermediate lobe, by means of the perifusion technique. Using our radioimmunoassay procedure, we found that serial dilutions of L. vivipara NIL extracts and synthetic alpha-MSH gave parallel binding curves. Administration of graded doses of TRH (10(-8)-10(-6) M) did not cause any modification of alpha-MSH release. In contrast, infusion of a depolarizing concentration of K+ induced a robust stimulation of alpha-MSH secretion. These results indicate that, in the lizard L. vivipara, the neuropeptide TRH does not stimulate pituitary melanotrophs.

Animals↗

Effects of corticosterone treatment on male aggressive behavior in a lizard (Anolis sagrei).

Social dominance in several species of lizards appears related to adrenal gland activity. The purpose of this study was to test whether the adrenal hormone corticosterone (CS) would inhibit the aggressive behavior of males of the lizard Anolis sagrei. Reproductively active males were randomly assigned to either CS or placebo (P) treatment groups or to an untreated stimulus male group. Both CS and P were administered in pellet form. Corticosterone pellets were designed to release approximately 4.8 micrograms of CS/day. Initially (Day 1), treatment males were implanted subcutaneously with a single pellet of either CS or P. After 1 week (Day 8), behavior tests were staged between treatment males and untreated stimulus males. The behavior of treatment males in response to stimulus males was videotaped. Significantly fewer CS-treated males than P-treated males approached the stimulus males, erected a crest, or showed biting behavior. Corticosterone-treated males also displayed significantly less frequently than did P-treated males and gave fewer displays of a type associated with dominance. Mean combined testis weight at the end of the experiment was significantly less in CS-treated males than in P-treated males and spermatogenic activity in CS-treated males was reduced. Plasma testosterone in CS-treated males was nondetectable, whereas P-treated males had normal levels of this steroid. These results indicate that CS can inhibit male aggressive behavior and testicular function in a lizard and support the hypothesis that the behavior changes observed in subordinate animals following defeat in agonistic encounters may in part be due to increased plasma CS levels.

Aggression↗

Interrenal activity in the female lizard Lacerta vivipara J.: in vitro response to ACTH 1-39 and to [Sar1, Val5] angiotensin II (ANG II).

A perifusion system technique was developed in order to determine in vitro the respective roles of ACTH and ANG II in the regulation of adrenal steroidogenesis in the lizard Lacerta vivipara. Synthetic human ACTH 1-39, administered as 20-min pulses, stimulated corticosterone (B) and aldosterone (A) release in a dose-dependent manner. The increase in corticosterone output was higher than that in aldosterone output, leading to an enhancement of the B/A ratio. Iterative stimulations with 1 nM ACTH (20-min pulses every 120 min) led to reproducible increases in corticosterone and aldosterone release. Prolonged stimulation with 1 nM ACTH (up to 240 min) caused a sustained increase in corticosteroid release, suggesting that, in the lizard, ACTH does not induce any desensitization phenomenon. The angiotensin II analogue [Sar1, Val5] ANG II also stimulated corticosterone and aldosterone release in a dose-dependent manner; the stimulatory effects of ANG II on both steroids were very similar. These results indicate that, in lizards, ACTH plays a major role in the regulation of adrenal steroidogenesis. Since ANG II stimulates the production of gluco- and mineralocorticoids, our data raise the question of the existence of two cell types synthesizing corticosterone and aldosterone, respectively, in reptiles.

Adrenal Glands↗