Patterns of olfactory projections in the desert iguana,Dipsosaurus dorsalis

The neural organization of the olfactory system in the desert iguana, Dipsosaurus dorsalis, has been investigated by using the Fink-Heimer technique to trace the efferents of the main and accessory olfactory bulbs, and Golgi preparations to determine the spatial relations between olfactory afferents and neurons in the primary olfactory centers. The accessory olfactory bulb projects to the ipsilateral nucleus sphericus via the accessory olfactory tract. The main olfactory bulb projects to the ipsilateral telen-cephalon via four tracts. The medial olfactory tract projects to the rostral continuation of medial cortex and to the septum. The intermediate olfactory tract projects to the olfactory tubercle and retrobulbar formation. The lateral olfactory tract projects to the rostral part of lateral cortex. The intermediate and lateral olfactory tracts also merge caudally to form the stria medullaris, which crosses the midline in the habenular commissure and distributes fibers to the contralateral hemisphere via two tracts. The lateral corticohabenular tract terminates in the contralateral lateral cortex. The anterior olfactohabenular tract terminates in the contralateral olfactory tubercle, retrobulbar formation and septum. The relation of olfactory afferents to neurons in the medial cortex, lateral cortex, nucleus sphericus, and septum corresponds to a pattern of organization that is typical of many olfactorecipient structures. Such structures are trilaminar, with neurons whose somata are situated in the intermediate layer (layer 2) sending spine-laden dendrites into an outer, molecular layer (layer 1). Olfactory afferents intersect the distal segments of these dendrites. By contrast, other olfactorecipient structures in Dipsoaurus deviate from the familiar pattern. Olfactory afferents intersect somata lying in layer 2 of the retrobulbar formation. Olfactory afferents include some fibers which course perpendicularly to the surface of the olfactory tubercle and extend deep to layer 2.     

Electrophysiological and behavioral correlates of sleep in the desert iguana, Dipsosaurus dorsalis Hallowell

The circadian behavior of the desert iguana, Dipsosaurus dorsalis, was investigated on the basis of behavioral observation and electrophysiological recording. D. dorsalis adequately complies with accepted criteria for both behavioral sleep and paradoxical sleep. At 20 degrees C, 12% of the photophase is spent in sleep, 95% of the scotophase is spent in sleep. Paradoxical sleep occurs at all times of the year, but only at temperatures of 20 and 30 degrees C. Amounts of behavioral sleep are affected by both temperature and time of year. Total sleep increases with decreased day length and decreased temperature. Daytime sleep increases with decreased temperature.     

Retinohypothalamic projections and immunocytochemical analysis of the suprachiasmatic region of the desert iguana Dipsosaurus dorsalis

Two separate and distinct retinal projections to the hypothalamus in the iguanid lizard Dipsosaurus dorsalis were described using horseradish peroxidase and cobalt-filling techniques. Both of the projections were unilateral and completely crossed; one terminated in the supraoptic nucleus and the other in the suprachiasmatic nucleus. Immunocytochemical analysis showed that the supraoptic nucleus contained cell bodies and fibers that cross-react with antibodies raised against arginine vasopressin, while the suprachiasmatic nucleus contained arginine vasopressin-like immunoreactive fibers emanating from cells in the nearby paraventricular nucleus. The suprachasmatic nucleus contained a dense plexus of fibers that cross-reacted with neuropeptide-Y antibody. Antiserum against vasoactive intestinal polypeptide showed no reactivity in any part of the forebrain, while antiserum against serotonin showed sparse and uniform reactivity throughout the forebrain, including the suprachiasmatic nucleus. These results, together with other data, indicate that the suprachiasmatic nucleus of D. dorsalis is homologous to the suprachiasmatic nuclei of rodents, structures known to contain circadian pacemakers. We suggest that the suprachiasmatic nucleus may play a similar role in the circadian system of D. dorsalis.     

Effect of activity duration on recovery and metabolic costs in the desert iguana (Dipsosaurus dorsalis).

The majority of elevated O(2) consumption associated with short and vigorous activity occurs during recovery, thus an assessment of associated metabolic costs should also examine the excess post-exercise oxygen consumption (EPOC). This study examined O(2) uptake during exercise, EPOC and distance traveled during 5-, 15-, 60- and 300-s sprints at maximal treadmill intensity in Dipsosaurus (N=10; 74.3+/-2.1 g). EPOC (0.08, 0.14, 0.23 and 0.18 ml O(2) g(-1), respectively) was large (80-99% of total elevated O(2) consumption) and increased significantly between 5 and 60 s. The cost of activity (C(act); ml O(2) g(-1) x km(-1)), intended to reflect the total net costs associated with the activity, was calculated as the total elevated O(2) consumption per unit distance traveled. C(act) decreased with activity duration due to proportionally larger increases in distance traveled relative to EPOC volume, and is predicted by the equation C(act)=14.7 x activity duration (s)(-0.24). The inclusion of EPOC costs provides an ecologically relevant estimate of the total metabolic cost of locomotor activity. C(act) exceeds standard transport costs at all durations examined due to the addition of obligate recovery costs. The differences are large enough to impact energy budget analyses for ectotherms.     

Metabolic support of moderate activity differs from patterns seen after extreme behavior in the desert iguana Dipsosaurus dorsalis

This study examined glucose and lactate metabolism in an iguanid lizard, Dipsosaurus dorsalis, during rest and after activity patterned on field behavior (15 s of running at 1 m/s). Metabolite oxidation and incorporation into glycogen by the whole animal, the liver, and oxidative and glycolytic muscle fibers were measured using (14)C- and (13)C-labeled compounds. Results showed that lactate metabolism is more responsive to changes that occurred between rest and recovery, whereas glucose appears to play a more steady state role. After activity, lactate oxidation produced 57 times as much ATP during 1 h of recovery than did glucose oxidation. However, lactate oxidation rates were elevated for only 30 min after activity, while glucose oxidation remained elevated beyond 1 h. Lactate was the primary source for glycogen synthesis during recovery, and glucose was the main glycogenic substrate during rest. This study supports previous research showing that brief activity in D. dorsalis is primarily supported by glycolysis and phosphocreatine breakdown, but it also suggests that there may be less of a reliance on glycolysis and a greater reliance on phosphocreatine than previously shown. The findings presented here indicate that the metabolic consequences of the behaviorally relevant activity studied are less severe than has been suggested by studies using more extreme activity patterns.     

Elevation of the panting threshold of the desert iguana,Dipsosaurus dorsalis,during dehydration: Potential roles of changes in plasma osmolality and body fluid volume

Summary Dehydration of the desert iguana,Dipsosaurus dorsalis, resulted in a progressive elevation in the magnitude of the skin temperature necessary to elicit thermal panting (i.e., the panting threshold). Panting threshold increased from 43.4±0.8 °C at 100% initial body weight (IBW) to 45.4±1.2 °C at 90% IBW to 45.7±0.9 °C at 80% IBW. Plasma osmolality showed no significant change with dehydration to 80% IBW. Changes in plasma osmolality, whether induced by NaCl or non-ionic sucrose loading, had a significant impact on panting threshold. Increasing plasma osmolality resulted in an elevation of panting threshold while decreasing plasma osmolality resulted in lower panting thresholds. Decreasing body fluid volume by exsanguination of 1 ml whole blood/100 g body weight resulted in a mean increase in panting threshold by 0.7±0.2 °C. Volume loading with 160 mM NaCl (approximately isosmotic) had no significant effect on panting threshold. These data suggest that plasma osmolality and decreases in body fluid volume may be potent modulators of panting threshold during periods of water deprivation. However, at least in desert iguanas, increases in plasma osmolality would not appear to be an important factor in the elevation of panting threshold during dehydration to 80% IBW.     

Contributions to elevated metabolism during recovery: dissecting the excess postexercise oxygen consumption (EPOC) in the desert iguana (Dipsosaurus dorsalis)

The excess postexercise oxygen consumption (EPOC), a measure of recovery costs, is known to be large in ectothermic vertebrates such as the desert iguana (Dipsosaurus dorsalis), especially after vigorous activity. To analyze the cause of these large recovery costs in a terrestrial ectotherm, Dipsosaurus were run for 15 s at maximal-intensity (distance 35.0+/-1.9 m; 2.33+/-0.13 m s(-1)) while O(2) uptake was monitored via open-flow respirometry. Muscle metabolites (adenylates, phosphocreatine, and lactate) were measured at rest and after 0, 3, 10, and 60 min of recovery. Cardiac and ventilatory activity during rest and recovery were measured, as were whole-body lactate and blood lactate, which were used to estimate total muscle activity. This vigorous activity was supported primarily by glycolysis (65%) and phosphocreatine hydrolysis (29%), with only a small contribution from aerobic metabolism (2.5%). Aerobic recovery lasted 43.8+/-4.6 min, and EPOC measured 0.166+/-0.025 mL O(2) g(-1). This was a large proportion (98%) of the total suprabasal metabolic cost of the activity to the animal. The various contributions to EPOC after this short but vigorous activity were quantified, and a majority of EPOC was accounted for. The two primary causes of EPOC were phosphocreatine repletion (32%-50%) and lactate glycogenesis (30%-47%). Four other components played smaller roles: ATP repletion (8%-13%), elevated ventilatory activity (2%), elevated cardiac activity (2%), and oxygen store resaturation (1%).     

Ion secretion by salt glands of desert iguanas (Dipsosaurus dorsalis)

Unlike the NaCl-secreting salt glands of many birds and reptiles, the nasal salt glands of lizards can secrete potassium as well as sodium, with either chloride or bicarbonate as the accompanying anion. The factors responsible for initiating secretion by the gland and the rates of cation and anion secretion were studied in the desert iguana, Dipsosaurus dorsalis. Lizards were given combinations of ions for several days, and secreted salt was collected daily and analyzed for sodium, potassium, chloride, and bicarbonate. Maximum total cation secretion rate was 4.4+/-0.38 micromol/g/d. Cation secretion ranged from 24% to 100% potassium; even high NaCl loads did not abolish potassium secretion. Maximum bicarbonate secretion was about 0.5 micromol/g/d; chloride was the predominant anion. Secretion rate increased only in response to those treatments that included potassium and/or chloride; sodium ions and other osmotic loads (e.g., sucrose) did not increase secretion. This is in contrast to birds and some other reptiles with salt glands, which initiate NaCl secretion in response to any osmotic load. The specificity of the response of the salt gland of Dipsosaurus may be related to the ecological importance of dietary potassium and chloride for herbivorous desert lizards.     

Phylogeographic histories of representative herpetofauna of the southwestern U.S.: mitochondrial DNA variation in the desert iguana (Dipsosaurus dorsalis) and the chuckwalla (Sauromalus obesus)

To determine whether genetic variation in representative reptiles of the southwestern U.S. may have been similarly molded by the geologic history of the lower Colorado River, we examined restriction site polymorphisms in the mitochondrial DNA (mtDNA) of desert iguanas (Dipsosaurus dorsalis) and chuckwallas (Sauromalus obesus). Observed phylogeographic structure in these lizards was compared to that reported for the desert tortoise (Xerobates agassizi), whose mtDNA phylogeny demonstrates a striking genetic break at the Colorado River. Both the desert iguana and chuckwalla exhibit extensive mtDNA polymorphism, with respective genotypic diversities G = 0.963 and 0.983, close to the maximum possible value of 1.0. Individual mtDNA clones, as well as clonal assemblages defined by specific levels of genetic divergence, showed pronounced geographic localization. Nonetheless, for each species the distributions of certain clones and most major clonal groupings encompass both sides of the Colorado River valley, and hence are clearly incongruent with the phylogeographic pattern of the desert tortoise. Overall, available molecular evidence provides no indication that the intraspecific phylogenies of the southwestern U.S. herpetofauna have been concordantly shaped by a singular vicariant factor of overriding significance.     

Intermittent locomotor activity that increases endurance also increases metabolic costs in the desert Iguana (Dipsosaurus dorsalis)

Intermittent activity, alternating bouts of activity and rest, can extend endurance relative to continuous locomotion. Utilizing a rapid fatiguing activity intensity (1.08 m s(-1)), Dipsosaurus dorsalis (n = 14) ran repeated bouts of varying durations (5, 15, or 30 s) interspersed with variable pause periods (100%, 200%, 400%, or 800% of the activity period) until exhausted. Total distance ran increased relative to continuous locomotion. The largest increases were seen when activity periods were limited to 5 s and pause periods were extended from 5 s to 20 s to 40 s (55, 118, and 193 m, respectively). To analyze these increases further, O(2) consumption was measured for six bouts of 5-s activity separated by either 5, 20, or 40 s (n = 8). The sum of elevated O(2) consumption during activity, pauses, and recovery increased significantly from 0.08 to 0.09 and 0.12 mL O(2) g(-1) as pause duration increased, primarily due to greater O(2) consumption during longer pause intervals. Postexercise recovery metabolism was a large cost (>57% of total) but did not differ among treatments. Overall, 40-s pauses were most expensive (absolutely and per unit distance) but provided the greatest endurance, likely due to further repletion of metabolites or removal of end products during the longer pause. In contrast, the shortest pause period was most economical but exhausted the animal most rapidly. Thus, a pattern of intermittent activity utilized by an animal may have energetic advantages that sometimes may be offset by behavioral costs associated with fatigue.     

Circadian locomotor rhythms in the desert iguana

Summary Desert iguanas, Dipsosaurus dorsalis, displaying freerunning circadian locomotor rhythms in conditions of constant darkness and temperature received electrolytic lesions to the hypothalamus. The locomotor activity of those lizards (N = 9) which sustained 80% or more damage to the suprachiasmatic nucleus (SCN) became arrhythmic whereas all animals that sustained less than 35% damage to the SCN remained rhythmic, even though they sustained significant damage to nearby regions of the hypothalamus and preoptic area. These results suggest strongly that the SCN plays a role in the regulation of circadian rhythms in the desert iguana. Taken together with other evidence, they support the view that this structure is homologous to the mammalian SCN, which acts as a pacemaker in the circadian system.Abbreviations SCN suprachiasmatic nucleus - freerunning circadian period     

Day-night variations in blood and intracellular pH in a lizard,Dipsosaurus dorsalis

Blood pH, PCO2 and PO2 of Dipsosaurus dorsalis were measured during the day and at night. Lizards at constant body temperature (25, 37 degrees C) and lizards experiencing diurnal changes in body temperature similar to those in nature were studied. In lizards at constant body temperatures, blood pH was about 0.1 unit less and blood PCO2 was 4-7 Torr higher at night compared to day. Similar patterns were seen in lizards on natural thermal cycles. Intracellular pH (pHi) of skeletal muscle, esophagus and liver was about 0.2 units lower at night than day but myocardial pHi was unchanged. Reduction in breathing frequency, and thus a relative hypercapnia from hypoventilation was consistent with the nocturnal acidification of the blood and intracellular compartments. Nocturnal acidification (CO2 retention) corresponds to periods of minimum metabolism. The possible impacts of diurnal shifts in hydrogen ion concentration on energy metabolism and metabolic regulation are discussed.     

Histochemical, enzymatic, and contractile properties of skeletal muscle fibers in the lizard Dipsosaurus dorsalis

Lizard skeletal muscle fiber types were investigated in the iliofibularis (IF) muscle of the desert iguana (Dipsosaurus dorsalis). Three fiber types were identified based on histochemical staining for myosin ATPase (mATPase), succinic dehydrogenase (SDH), and alphaglycerophosphate dehydrogenase (alphaGPDH) activity. The pale region of the IF contains exclusively fast-twitch-glycolytic (FG) fibers, which stain dark for mATPase and alphaGPDH, light SDH. The red region of the IF contains fast-twitch-oxidative-glycolytic (FOG) fibers, which stain dark for all three enzymes, and tonic fibers, which stain light for mATPase, dark for SDH, and moderate for alphaGPDH. Enzymatic activities of myofibrillar ATPase, citrate synthase, and alphaGPDH confirm these histochemical interpretations. Lizard FG and FOG fibers possess twitch contraction times and resistance to fatigue comparable to analogous fibers in mammals, but are one-half as oxidative and several times as glycolytic as analogous fibers in rats. Lizard tonic fibers demonstrate the acetylcholine sensitivity common to other vertebrate tonic fibers.