Identity and distribution of Agama stellio picea Parker (Sauria: Agamidae), endemic to the volcanic desert of Jordan

Abstract

Agama stellio picea Parker, 1935, was clearly defined as a black lizard, with the female dotted with orange, from the Transjordan lava desert. Three later collections confirmed this. While the subspecies appears to be endemic to Jordan, its name has been erroneously applied to other A. stellio populations in adjacent countries.     

Purification and kinetic properties of skeletal muscle lactate dehydrogenase from the lizard Agama stellio stellio

Lactate dehydrogenase isoenzyme LDH-5 (M4) was purified to homogeneity from the skeletal muscle of lizard Agama stellio stellio as a poikilothermic animal, using colchicine-Sepharose chromatography and heat inactivation. The purified enzyme showed a single band after SDS-PAGE, corresponding to a molecular weight of 36 kD. The K _m values for pyruvate, NADH, lactate, and NAD⁺ were 0.020, 0.040, 8.1, and 0.02 mM, respectively. Pyruvate showed maximum activity at about 180 M, with a decline at higher concentrations. The enzyme was stable at 70°C for 30 min, but was rapidly inactivated at 90°C. The optimum pH for the forward reaction (pyruvate to lactate) was 7.5, and for the reverse reaction (lactate to pyruvate) was 9.2. Oxalate, glutamate, Cu²⁺, Co²⁺, Mn²⁺, and Mg²⁺ were inhibitory in both forward and reverse reactions.     

Genetic variation and climatic selection in the lizard Agama stellio in Israel and Sinai

Summary Allozymic variation in proteins encoded by 25 loci was analyzed electrophoretically in 242 adult specimens representing nine populations of the Levantine lizard, Agama stellio, comprising two subspecies: the Mediterranean A. stellio subsp., and the desert-inhabiting A. stellio brachydactyla from the Negev and Sinai. Likewise, four body traits were measured in the same populations. The nine populations were sampled along a general southward transect of increasing aridity. Agama stellio is above average in both polymorphism, P, and heterozygosity, H, as compared to other reptiles and vertebrates in general, displaying levels of genetic variation characterizing habitat generalist vertebrates. In the populations studied no fixation of alternative alleles was found in any of the 25 loci: rather the commonest allele was either fixed or predominated in 23 of 25 loci examined. Eleven loci (44%) were monomorphic in all nine populations. However, of the remaining 14 polymorphic loci, eight were strongly polymorphic displaying distinct genetic differentiation between populations. Genetic diversity (indexed by P and H) displayed geographic variation and was slightly higher in A.s. brachydactyla than in A. stellio subsp. Nevertheless, genic similarity between populations was high. A statistically significant amount of morphological variation between localities was found for all body characters. In general, body size increased southwards and eastwards with aridity.Selection at some loci is suggested by significant deviation from Hardy-Weinberg expectations and possibly by excess heterogeneity of effective inbreeding coefficients, F_e. Furthermore, allozymic variation at seven loci (Ldh-1, Idh-1, 6-Pgd-1, Aat-1, Pgm-2, Pept-1, and Trf) and geographic variation in body size and weight were significantly correlated with, and predictable by, climatic variables, primarily by water availability and secondarily by temperature. Finally, allozymic and morphological variations were partly correlated.The spatial patterns and ecological correlates of genic and morphological variations in Agama stellio in Israel and Sinai suggest that at least some proteins and body size differentiate geographically and appear to be adaptive, presumably with respect to factors affecting the availability of water.     

Prey capture in the lizard Agama stellio

Prey capture in Agama stellio was recorded by high-speed video in combination with the electrical activity of both jaw and hyolingual muscles. Quantification of kinematics and muscle activity patterns facilitated their correlation during kinematic phases. Changes in angular velocity of the gape let the strike be subdivided into four kinematic phases: slow open (SOI and SOII), fast open (FO), fast close (FC), and slow close-power stroke (SC/PS). The SOI phase is marked by initial activity in the tongue protractor, the hyoid protractor, and the ring muscle. These muscles project the tongue beyond the anterior margin of the jaw. During the SOII phase, a low level of activity in the jaw closers correlates with a decline of the jaw-opening velocity. Next, bilateral activity in the jaw openers defines the start of the FO phase. This activity ends at maximal gape. Simultaneously, the hyoid retractor and the hyoglossus become active, causing tongue retraction during the FO phase. At maximal gape, the jaw closers contract simultaneously, initiating the FC phase. After a short pause, they contract again and the prey is crushed during the SC/PS phase. Our results support the hypothesis of tongue projection in agamids by Smith ([1988] J. Morphol. 196:157–171), and show some striking similarities with muscle activity patterns during the strike in chameleons (Wainwright and Bennett [1992a] J. Exp. Biol. 168:1–21). Differences are in the activation pattern of the hyoglossus. The agamid tongue projection mechanism appears to be an ideal mechanical precursor for the ballistic tongue projection mechanism of chameleonids; the key derived feature in the chameleon tongue projection mechanism most likely lies in the changed motor pattern controlling the hyoglossus muscle. © 1995 Wiley-Liss, Inc.     

On the distribution of four species of agama (Agamidae) in Turkey

Abstract

The distribution of the Hardun, Agama stellio, the Caucasian Agama, Agama caucasia, the Syrian Agama, Agama ruderata, and Phrynocephalus helioscopus, in Turkey is presented with dot maps A. stellio and A. caucasia are sharply separated both horizontally and vertically. The area of A. stellio is defined by the March isotherm of 8°C and the July and August isotherms of 24°C. A. ruderata occurs in the large steppe areas of Inner and South-east Anatolia, but not in those of Eastern Anatolia.     

Purification and characterization of cathepsin L from skeletal muscle of the lizard Agama stellio stellio

Cathepsin L from skeletal muscle of the lizard Agama stellio stellio was purified to homogeneity by ion-exchange and gel-permeation chromatography. The molecular weight of the cathepsin L is estimated to be 34 kD, and its isoelectric point is 5.5. The cathepsin L has a pH optimum of 6.1, requires a thiol-reducing reagent for activation, and is inhibited by cysteine protease inhibitors. The Km and kcat values for Z-Phe-Arg-MCA as substrate are 1.4 microM and 6.2 sec-1, respectively. This enzyme readily hydrolyzes proteins such as insulin B chain, hemoglobin, and serum albumin.     

Activity metabolism of the lizard Agama stellio stellio

Oxygen consumption and lactate production above resting levels, and selected body temperatures, were measured in the lizard Agama stellio. Active and resting VO2 have low Q10 (1.7, 2.0) in the activity range 30-37 degrees C and higher Q10 (3.8, 4.0) below this. A correlation was found between published resting and active VO2 of lizards, and between VO2 and lifestyle. Four types were recognized, in order of increasing VO2: (a) fossorial; (b) sit-and-wait (including A. stellio); (c) cruising, and (d) widely foraging. A. stellio has a high capacity for lactate production, correlated with its short but rapid bursts of activity. This accounts for 80-90% of the energy used during 30 sec maximal activity.     

Fibre types in the locomotory muscles of an antarctic teleost,Notothenia rossii

Summary The metabolic and structural differentiation of locomotory muscles of Notothenia rossii has been investigated. In this species sustained locomotion is achieved by sculling with enlarged pectoral fins (labriform locomotion), whilst the segmental myotomal muscle is reserved for burst activity. Red, white and subepidermal fibres can be distinguished in the trunk by histochemical and ultrastructural criteria. The main pectoral muscle (m. adductor profundus) consists entirely of red fibres. These three main fibres types show differences in histochemical staining profiles, capillarization, myofibril shape and packing, and lipid and mitochondrial content. The fractional volume of mitochondria amounts to 38% for pectoral, 30% for red myotomal and 1.9% for white myotomal fibres. Enzyme activities of red pectoral muscle are consistent with a higher potential for aerobic glucose and fatty acid oxidation than for the red myotomal fibres. Mg²⁺ Ca²⁺ -myofibrillar ATPase activities are similar for red pectoral and myotomal muscles and approximately half of those white fibres. Specialisations of N. rossii muscles associated with labriform swimming and locomotion at Antarctic temperatures are discussed.     

Structure and histochemical organization of the spleen of Agama stellio (Sauria: Agamidae) and Chalcides ocellatus (Sauria: Scincidae)

The spleen of Agama stellio is composed mainly of red pulp; the white pulp is poorly developed, and its clusters are scattered throughout the organ and contain lymphocytes, reticular cells, and some plasma cells. The red pulp consists of clear reticular cells intermingled with blood cells, sinusoids, and pigment cells. The spleen of Chalcides ocellatus is encapsulated by connective tissue and is composed of white and red pulp. The white pulp consists of lymphoid tissue that surrounds the central arterioles, forming the periarteriolar lymphocyte sheath (PALS). The red pulp is composed of a system of venous sinuses and cords. The results of various histochemical procedures designed to demonstrate mucosubstances, proteins, and nucleic acids indicate that the spleen in these species resembles the mammalian spleen. © 1993 Wiley-Liss, Inc.     

Isolation, purification, and properties of aminopeptidase H from skeletal muscle of the lizard Agama stellio stellio

Aminopeptidase H was isolated and purified from fresh skeletal muscle of the lizard Agama stellio stellio by ammonium sulfate fractionation and successive chromatographies on DEAE-cellulose, Ultrogel AcA-34, activated thiol-Sepharose 4B, phenyl-Sepharose CL-4B, and DEAE-cellulose again. This is the first report of the isolation of aminopeptidase H from a reptile. The purified enzyme migrated as a single band on SDS-PAGE. The molecular weight of the enzyme was 48 kD by SDS-PAGE and 384 kD on Ultrogel AcA-34 column chromatography. The optimum pH for hydrolysis of L-leucine beta-naphthylamide (Leu-Nap) was 7.8. The Km values for the hydrolysis of Leu-Nap and Nalpha-benzoyl-DL-arginine beta-naphthylamide (BzArg-Nap) were 0.48 and 0.99 mM, respectively. These activities were strongly inhibited by iodoacetic acid and leupeptin but were not affected by EDTA, pepstatin, bestatin, or phenylmethylsulfonyl fluoride. The enzyme has been shown not to hydrolyze proteins such as hemoglobin, BSA, myofibrillar proteins, and sarcoplasmic proteins.     

Diet and reproductive biology of the Starred Agama,Laudakia stellio (Linnaeus, 1758) (Squamata: Agamidae), in the northern Sinai,Egypt

We quantified sexual size dimorphism, diet and reproduction in the Starred Agama, Laudakia stellio, in northern Sinai. Males were larger than females in snout-vent length and head index. The species is a sit-and-wait predator and feeds on insects, mainly coleopterans. About 30% of stomachs included plant material. No difference between sexes existed in terms of prey size preference. The reproductive season is seen to be year round with no distinctive seasonality. The smallest sexually mature female measured 92 mm SVL, whereas the smallest sexually reproductive male was 89 mm SVL. Clutch size ranged from 6 to 18 eggs.     

Expression of calcium channels in Xenopus oocyte injected with crab skeletal muscle fibre mRNA

Summary— Using the whole cell voltage-clamp technique and a Cl free and Na free Ba methane sulfonate solution, stage V and VI Xenopus oocytes demonstrated a Ba current (endogenous component) with a peak amplitude average of 6 nA (6 ± 2 nA). When oocytes were injected with crustacean skeletal muscle mRNA, an additional component of I_Ba could be detected (exogenous I_Ba). The latter current could be distinguished from the native one by several electrophysiological means: a peak amplitude average of 90 nA (90 ± 4 nA), activation potential threshold, steady state inactivation properties and sensitivity to Ca blockers. As shown by Jdaïâa and Guilbault in crustacean skeletal muscle fibres, exogenous I_Ba could be divided into two components: a “fast component” and a “slow component” probably passing through two types of Ca channels (fast and slow) since the peak Ba current voltage relationship was biphasic and the fast component of exogenous I_Ba was less sensitive than the slow to nifedipine. The features of the newly synthesized channels incorporated in the Xenopus oocyte membrane suggest that they may be associated with fast and slow channels, previously described in many preparations, particularly in crustacean skeletal muscle fibres.     

Myotube production in fast myotomal muscle is switched-off at shorter body lengths in male than female Atlantic halibut Hippoglossus hippoglossus (L.) resulting in a lower final fibre number

A sampling method is described to determine accurately the number of fast myotomal muscle fibres (N_F) in a large flatfish species, the Atlantic halibut Hippoglossus hippoglossus. An unusual feature of the fast myotomal muscle is the presence of internalized strips of slow muscle fibres. In fish of 1·5–3·5 kg (n = 24), the total cross‐sectional area (A_TC) of fast muscle was 18% greater in the dorsal than ventral myotomal compartments (P < 0·05), whereas there was no significant difference between left‐ and right‐hand sides of the body. Due the bilateral asymmetry, muscle blocks (5 × 5 × 5 mm) were prepared to systematically sample each myotomal quadrant (dorsal, ventral, left‐ and right‐side) and the diameters of 150 fast fibres measured per block. Smooth non‐parametric probability functions were fitted to a minimum of 800 measurements of fibre diameter per quadrant (n = 5). There were no significant differences in the distribution of muscle fibre diameters between myotomal compartments and therefore N_F could be estimated from a single quadrant. The number of blocks required to estimate N_F with a repeatability of ±2·5% increased from six at 300 g body mass to 17 at 96·5 kg, caused by variation within and between blocks. Gompertz curves were fitted to measurements of fibre number and fork length (L_F). The estimated final fibre number was 8·96 × 10⁵ (7·99–9·94 × 10⁵, 95% CI) for males and 1·73 × 10⁶ (1·56–1·90 × 10⁶, 95% CI) for female fish. The estimated L_F for cessation of fibre recruitment in the fast muscle of female fish (1775 mm) was almost twice that in males (810 mm), reflecting their greater ultimate body size.     

Actomyosin ATPase activity in Atlantic hagfish muscles

Summary Incubation for Ca⁺⁺-activated myosin ATPase reveals three types of muscle fibres in m. parietalis of the Atlantic hagfish (Myxine glutinosa), while m. craniovelaris and m. longitudinalis linguae both contain one type of muscle fibres.The fast twitch white fibres of m. longitudinalis linguae and m. parietalis show relatively high ATPase activity, while the intermediate fibres of m. parietalis show low activity. Despite of being slow non-twitch, the superficial red fibres of m. parietalis and the fibres of m. craniovelaris show an ATPase activity even higher than that of the fast twitch muscle fibres.     

The physiological basis of slow locomotion in chamaeleons

The African chamaeleon, Chamaeleo senegalensis, will not move faster than approximately 0.1 m/second at 23 degrees C, whereas the lizard Agama agama, like most lizards its size, runs at speeds more than 10X as fast. To account for this difference, we measured various physiological parameters of the iliofibularis muscle of both lizards. The maximum speed of tetanic contraction of unloaded Chamaeleo muscle was half as fast as that of Agama muscle (2.5 vs. 5.8 resting lengths per second). Heavily loaded Chamaeleo iliofibularis contracted at nearly 1/4 the speed of Agama muscle. Time to peak isometric twitch tension and time to half relaxation were twice as long in Chamaeleo as in Agama (122 vs. 58 msec, and 168 vs. 81 msec). Much more of the Chamaeleo muscle consisted of tonic muscle fibers, and the Chamaeleo muscle, compared to Agama muscle, showed physiological evidence of having a significant amount of tonic fibers (potassium contracture and high tetanus to twitch ratios). Finally, the myofibrillar ATPase activity of the Chamaeleo muscle was 1/3 that of Agama muscle. Thus, these results show that the slow locomotion of old world chamaeleons can, in part, be explained by the physiology, biochemistry, and fiber-type distribution of their muscles.     

Muscular waves contribute to gliding rate in the freshwater gastropod Lymnaea stagnalis

This study revises the mechanisms of ciliary locomotion and demonstrates muscular contribution to locomotion rate in Lymnaea stagnalis. L. stagnalis sticks to the substratum by the foot sole and moves smoothly with no visible contractions of the foot. A ciliated epithelium covering the sole is underlain by smooth muscle cells containing giant mitochondria. It is shown here that slow (basal) locomotor activity (measured as the flow rate of physiological saline over isolated sole) appears spontaneously or is induced by 10⁻⁸–10⁻⁷ M 5-HT. 5-HT (10⁻⁷–10⁻⁴ M) facilitates locomotor activity dose-dependently, and KCN (an inhibitor of mitochondrial respiration) decreases these effects to the basal level. 5-HT and KCN have no effect on the frequency of ciliary beat (stroboscopic measurements), and blockers of anaerobic glycolysis inhibit ciliary motility. Under anaerobic conditions locomotion of a snail is slow and insensitive to 5-HT in contrast to that in aerobic environments. It is concluded that glycolysis supplies energy to ciliated cells and respiration to sole muscle cells; 5-HT stimulates ciliary beating in an all-or-none fashion and muscular waves in a dose-dependent manner; cilia provide slow (basal) gliding, and locomotory rate up to 80% above the basal level is determined by muscular waves.     

Sexual dimorphism in histological characteristics and contractility of the iliofibularis muscle in the lizard Sceloporus torquatus

The iliofibularis is a hindlimb muscle used in lizard locomotion that is composed of at least three types of fibres: fast‐twitch‐glycolytic (FG), fast‐twitch‐oxidative‐glycolytic fibre (FOG) and slow‐twitch‐oxidative (SO). The striated skeletal muscle is a highly plastic tissue undergoing phenotypic change in response to activity. The lizard Sceloporus torquatus has sexual differences associated with microhabitat use, which can be reflected in the physiology and anatomy of the muscle, and thus, in our study, we analysed the morphological and contractile characteristics of the iliofibularis muscle (IF) of S. torquatus males and females. We found a larger prevalence of FOG compared with FG and SO fibres in the muscle of both sexes. We also found that males show larger areas of the three types of fibres, develop greater strength but also faster fatigue than females, suggesting that strength is a key functional feature that enables males to perform faster movements (but for shorter periods), associated with the demands of territoriality.     

Ultrastructure and metabolism of skeletal muscle fibres in the tench: Effects of long-term acclimation to hypoxia

Summary Tench (Tinca tinca) were acclimated to either aerated (P _{O 2} 17.6 KPa) or hypoxic (P _{O 2} 1.5 KPa) water for 6 weeks.Acclimation to hypoxia resulted in a decrease in mitochondrial volume fraction in both slow (22.9 to 15.0 %) and fast glycolytic (4.5 to 1.8 %) myotomal muscles fibres (P<0.01).Intermyofibrillar mitochondrial populations (4.4 to 1.2% slow; 0.6 to 0.04% fast fibres) were affected to a greater extent than those in the subsarcolemmal zone (18.5 to 13.8% slow; 3.9 to 1.8% fast fibres). After acclimation to hypoxia, cytochrome-oxidase activities decreased by 31 and 33 % in slow and fast fibres, respectively, but were maintained in the liver.Fibre size remained unchanged and actively differentiating fibres were observed in muscles from both groups of fish. Hypoxia resulted in a significant increase in myofibrillar volume fraction in both slow (43.1 to 56.1 %) and fast glycolytic fibres (73.1 to 82.7%) (P<0.05).Glycogen concentrations (mg/100g tissue) for liver (6616) slow muscle (1892) and fast muscle (334) were similar for fish acclimated to aerated or hypoxic water. Acclimation to hypoxia increased carnitine palmitoyl transferase activity (moles substrate utilised g·dry wt_-1 min^-1) in slow (0.42 to 1.1), fast glycolytic muscle (<0.01 to 0.15) and liver (1.1 to 3.7) indicating an enhanced capacity for fatty acid oxidation.Phosphofructokinase activities of fast glycolytic fibres were similar in fish acclimated to either aerated or hypoxic water, consistent with an unaltered capacity for anaerobic glycogenolysis. Hexokinase activities (moles substate utilised, g·dry wt^-1 min^-1) decreased in fast fibres (1.2 to 0.4) but were maintained in the slow muslce (2.1 to 2.5) and liver (4.5 to 4.8) of hypoxic fish. The activities of phosphofructokinase in slow muscle and phosphofructokinase, pyruvate kinase and lactate dehydrogenase in liver were two times higher in fish acclimated to hypoxia. An enhanced capacity for glycolysis in these tissues may reflect a reduced threshold for anaerobic metabolism during activity and/or an adaptation for acute exposure to anoxia in fish acclimated to hypoxia.Abbreviations/Glossary CO cytochrome oxidase activity - CPT carnitine palmitoyltransferase activity - HK hexokinase activity - LDH lactate dehydrogenase activity - PFK phosphofructokinase activity - PK pyruvate kinase activity - Vv volume fractions of cell components - normoxic fish acclimated to aerated water - hypoxic fish acclimated to reduced oxygen tensions - P _{O 2} partial pressure of oxygen tension A preliminary account of part of this work was presented at theXth European Meeting on Muscle and Cell Motility held at Galway, Ireland, in September 1981     

The effect of elevated levels of thyroxine on the aerobic capacity of locomotor muscles of the tufted duck,Aythya fuligula

Following extended periods of relative inactivity, or prior to migration, birds are able to increase the aerobic capacity of their locomotory muscles. Thyroid hormones may influence this process. A preliminary study was undertaken to assess the ability of elevated levels of thyroxine to increase the aerobic capacity of the locomotory and cardiac muscles of adult tufted ducks. Administration of thyroxine in the food for 8 weeks had little effect on body mass or on the masses of the pectoralis, semitendinosus and iliofibularis muscles, although there were increases in resting oxygen consumption and in the mass of the cardiac ventricles. The maximum activity of the aerobic enzyme, citrate synthase, was significantly greater in the left ventricle, liver, and iliofibularis muscles (P<0.005) of treated birds. However, while there was clearly no difference in activity in the semimembranosus leg muscle, that of the pectoralis was not quite significant (P=0.078). It is concluded that addition of supra-physiological levels of exogenous thyroxine may induce a differential increase in the maximum activity of citrate synthase in the locomotor muscles of the tufted duck, which is correlated with the fibre type composition of these muscles. These results are consistent with those found in studies on rats, with slow oxidative fibres being the most sensitive, and fast glycolytic fibres the least sensitive, to thyroxine treatment.Abbreviations BM body mass - CS citrate synthase - CYTOX cytochrome c oxidase - FG last glycolytic - FOG fast oxydative glycolytic - VO₂ oxygen consumption - SO slow oxidative - T₄ thyroxine - T₃ triiodothyronine     

Fine structure and metabolism of multiply innervated fast muscle fibres in teleost fish

Summary Both the fast and slow muscle fibres of advanced teleost fish are multiply innervated. The fraction of slow-fibre volume occupied by mitochondria is 31.3%, 25.5% and 24.6%, respectively, for the myotomal muscles of brook trout (Salvelinus fontinalis), crucian carp (Carassius carassius), and plaice (Pleuronectes platessa), respectively. The corresponding figures for the fast muscles of these species are 9.3%, 4.6% and 2.0%, respectively. Cytochrome-oxidase and citrate-synthetase activities in the fast muscles of 9 species of teleost range from 0.20–0.93 moles substrate utilised, g wet weight muscle^-1 min^-1 (at 15° C) or around 4–17% of that of the corresponding slow fibres. Ultrastructural analyses reveal a marked heterogeneity within the fast-fibre population. For example, the fraction of fibres with <1% or >10% mitochondria is 0,4,42% and 36, 12 and 0%, respectively, for trout, carp and plaice. In general, small fibres (<500 m²) have the highest and large fibres (>1,500 m²) the lowest mitochondrial densities. The complexity of mitochondrial cristae is reduced in fast compared to slow fibres.Hexokinase activities range from 0.4–2.5 in slow and from 0.08–0.7 moles, g wet weight^-1 min^-1 in fast muscles, indicating a wide variation in their capacity for aerobic glucose utilisation. Phosphofructokinase activities are 1.2 to 3.6 times higher in fast than slow muscles indicating a greater glycolytic potential. Lactate dehydrogenase activities are not correlated with either the predicted anaerobic scopes for activity or the anoxic tolerances of the species studied. The results indicate a considerable variation in the aerobic capacities and principal fuels supporting activity among the fast muscles of different species. Brook trout and crucian carp are known to recruit fast fibres at low swimming speeds. For these species the aerobic potential of the fast muscle is probably sufficient to meet the energy requirements of slow swimming.