Blood Viscosity and Hematocrit in the Estuarine Crocodile, Crocodylus porosus
|
|
- Claud Atkins
- 6 years ago
- Views:
Transcription
1 Comparative Biochemistry and Physiology Part A: Physiology (1991) 99 (3): Blood Viscosity and Hematocrit in the Estuarine Crocodile, Crocodylus porosus R. M. G. Wells, L. A. Beard and G. C. Grigg Abstract 1. Viscosities of whole blood and plasma from Crocodylus porosus were low in comparison with other diving animals when measured over a range of shear rates at 30 C (e.g ± 0.07 mpa.sec and 1.67 ± 0.06 mpa.sec respectively, at a shear rate of 450 sec-1). 2. Hematocrit (19.2 ± 0.5%) and mean cell hemoglobin concentration (262 ± 23 g/l) are high in crocodiles compared with non-diving reptiles. 3. Nucleation of the red blood cell does not seem to have affected viscosity. 4. The potential for oxygen transport was estimated from pooled blood reconstituted to a range of hematocrits and used to predict an optimal hematocrit of 38%, which greatly exceeded measured values. 5. Low blood viscosity is associated with low flow resistance and may assist circulation during the low blood pressure events occurring during diving. Introduction Reptiles make extensive use of anaerobic metabolism during activity (Gatten, 1985), yet the natural dives of alligators (Lewis and Gatten, 1985) and crocodiles (Grigg et al., 1985) appear essentially aerobic. Crocodiles have a number of features that appear adaptive in their aquatic lifestyle and diving behaviour. These include cardiovascular shunts that conserve oxygen during diving (Grigg and Johansen, 1987), low respiratory quotients (Grigg, 1978), and hematocrits near the upper range of oxygen carrying capacity for reptiles (Grigg and Cairncross, 1980). Periods of apnoea have important consequences for blood gas tensions and acid base balance in crocodiles and, following burst exercise, result in an extensive lactacidosis (Seymour et al., 1985). Thus a store of oxygen and distribution to tissues is necessary for maintaining aerobic operations. There is, however, a potential conflict. High hematocrit pro-motes an increased blood oxygen store but it may be counter-productive to oxygen delivery if it also leads to higher viscosity. Blood viscosity and its effects on blood flow have been studied in a range of vertebrates, including mammals (Chien et al., 1971), amphibians (Hillman et al., 1985), and fishes (Fletcher and Haedrich, 1987; Wells and Forster, 1989), and these studies confirm that hematocrit has an exponential effect on blood viscosity. Further, diving mammals have very high blood viscosity, having high hematocrits, prompting Hedrick et al. (1986) to suggest that oxygen transport may become limited by blood viscosity. This possibility arises because blood oxygen transport is dependent on both blood flow and oxygen carrying capacity and, above a defined oxygen capacity, the former declines due to the blood's resistance to flow (Crowell and Smith, 1967). Nevertheless, in diving mammals, hematocrit is commonly elevated beyond the optimum predicted by this model (Hedrick et al., 1986; Wickham et al., 1989).
2 Apparently there are no data on the rheological characteristics of reptiles, despite the fact that all the orders have representatives that are competent diversmarine iguanas, aquatic snakes, turtles, and crocodiles. In this study, we examined the influence of hematocrit on blood flow, as revealed by viscosity measurements in the estuarine crocodile, Crocodylus porosus. Blood viscosity is a complex variable which is influenced by physical and physiological factors such as the deformability of erythrocytes, shear rate, and plasma proteins, and these also have been considered. Materials and Methods Blood sampling Estuarine crocodiles (Crocodylus porosus), 3-5 years old, and approximately 1.5 m in length were obtained during commercial slaughter at the Edward River Crocodile Farm at Pormpuraaw, Cape York Peninsula, Australia (14 54'S, 'E). Resting animals were killed with a rifle shot to the hind brain using low velocity 0.22 ammunition. Blood was taken immediately from a cerebral artery into a heparinised syringe and studied without delay. Hematology Hematocrit (Hct) and hemoglobin concentration ([Hb]) were measured using standard techniques (Dacie and Lewis, 1984), with the added precaution of centrifuging the cyanmethemoglobin solutions (Gruca and Grigg, 1980). Mean cell hemoglobin concentration (MCHC) was calculated from [Hb]/Hct fraction. Total plasma protein was determined spectrophotometrically by the biuret reaction. Blood viscosity Viscosity was measured on 0.5 ml samples of whole blood and separated plasma using a cone-plate viscometer with cone angle of 8 (model LVTD CP/11, Brookfield Engineering Labs, U.S.A.). Viscosity determinations were made over a range of shear rates corresponding with the rotational speed of the cone, and the results reported in units of mpa.sec (= centipoise). The temperature of the sample cup was held constant at C. Cells and plasma were separated by centrifugation at 1200 g from pooled blood, and a range of hematocrits reconstituted for subsequent viscometry. No evidence of lysis or change in MCHC occurred during the procedure. Fig. 1. Viscometric curves for blood and plasma from C. porosus at 30 C, Mean ± SD for seven animals.
3 Results Blood from C. porosus decreased in viscosity when shear rate was increased (Fig. 1) as expected from the non-newtonian behaviour of mammalian blood (Rand et al., 1964). Plasma viscosity was relatively less shear-dependent over the range of shear rates measured. Table 1 summarises hematocrit measurements and other hematological data. Hematocrit influenced blood viscosity in reconstituted samples and viscosity rose dramatically at hematocrits above 20%, this value corresponding with the in vivo hematocrit of 19.2% (Fig. 2). Assuming that oxygen transport by the blood is directly proportional to viscosity (see Snyder, 1983), the potential transport capacity may be calculated by the ratio 1.34 [Hb]/η, where η is viscosity at a shear rate of 90 sec 1 (Hedrick et al., 1986). The potential oxygen transport capacity for reconstituted hematocrits is shown in Fig. 3, and indicates an optimal hematocrit of about 38%. Fig. 2. Dependence of viscosity on hematocrit for reconstituted, pooled blood. Discussion High hematocrit and high MCHC combine to produce a high oxygen carrying capacity in C. porosus. Hematocrit values were slightly lower in our captive crocodiles than those in the wild (Grigg and Cairncross, 1980), and lower than in sea turtles (Lutz and Bentley, 1985), yet consistent with their diving habits, they lie in the upper range of oxygen carrying capacity for reptiles (Dawson and Poulson, 1962). Hematocrit comparisons in reptiles are however, complicated by differences in the temperatures at which the reptiles are normally active, such that hematocrit may be limited by the expected rise in blood viscosity as temperature falls (Pough, 1980). Values of MCHC were also high, consistent with the concept that high values of MCHC are typical of diving reptiles, compared with non-divers (Wood and Johansen, 1974; Seymour et al., 1981; Maginniss et al., 1983; Birchard et al., 1984), as is also the case within the Mammalia (Kooyman et al., 1981). In sea turtles also, pulmonary oxygen stores are supplemented by high blood oxygen carrying capacity and appear adequate to sustain predominantly aerobic dives (Lutz and Bentley, 1985). Diving in the estuarine crocodile is also predominantly aerobic (Grigg et al., 1985). Our results
4 therefore support the view that MCHC is a means of increasing oxygen stores in diving reptiles. The reptilian vascular system is characterised by large, widely spaced capillaries (Pough, 1980). Since reptilian hematocrits are only about one-third those of diving mammals, one might expect circulation under low pressure with little peripheral vasodilation to perfuse the predominantly white, skeletal musculature. The viscosity of crocodile blood at a high shear rate is slightly higher than that of diving mammals at an equivalent hematocrit of 20% (cf. Hedrick et a!., 1986; Wickham et al., 1989), but allowing for the 7 C higher temperature of the latter, they are most probably identical. This suggests that MCHC and nucleated erythrocytes in the crocodile do not impact significantly on blood viscosity. Increasing MCHC is an obvious way of increasing oxygen carrying capacity of the blood without increasing the number of circulating erythrocytes and hence viscosity. A marked increase in viscosity was evident above 20% hematocrit (Fig. 2). Fig. 3. Oxygen transport capacity as a function of hematocrit. In the case of C. porosus, the predicted hematocrit is far in excess of measured values. The theoretical optimum hematocrit for the elephant seal was lower than that measured in vivo, and has led to the suggestion that oxygen transport is compromised by storage capacity in diving mammals (Hedrick et al., 1986). Among poikilothermic divers, however, bull-frogs maintain a hematocrit that maximises oxygen transport capability (Weathers, 1976), but during maximal rates of oxygen uptake when
5 hematocrit is artificially elevated, the rise in viscosity cannot be compensated (Hillman et al., 1985). We propose that the lower than "optimal" hematocrit could be interpreted in terms of cardiovascular adjustments correlated with diving and other breathholds. Crocodiles are periodic breathers, and the pulmonary oxygen store is tapped during breathholds (Glass and Johansen, 1979); during longer dives blood may be shunted away from the lungs, perhaps in order to extend the aerobic range (Grigg and Johansen, 1987). Extended use of the pulmonary oxygen store is facilitated not only by the high blood oxygen carrying capacity, but by a significant increase in affinity of blood when carbon dioxide is given off from blood to lungs (Jelkmann and Bauer, 1980). The bicarbonate-mediated decrease in affinity ensures that oxygen released from the blood occurs at relatively high oxygen pressures. The oxygen binding properties of crocodile blood have been interpreted as being consistent with short burst activity (Grigg and Gruca, 1979), rather than for sustained aerobic activities such as diving. The oxygen affinity is relatively high, and there is a large effect of carbon dioxide (Glass and Johansen, 1979). The apparent conflict between the oxygen stores required for diving and the transport requirement for activity may be resolved in crocodiles by elevated reptilian MCHC and hematocrit that nevertheless do not compromise blood flow through high viscosity. The peculiar anatomy of the cardiovascular system in C. porosus allows for a pulmonary by-pass shunt during diving, and blood pressure decreases (Grigg, 1989). This arrangement also allows for extended aerobic metabolism by diverting blood to the systemic system. A low blood viscosity may assist circulation over a range of reduced blood pressures associated with diving. Acknowledgements We thank the staff of the Edward River Crocodile Farm and, in particular, Don Morris and Vic Onions for facilitating our research. The authors are grateful to the Australian Research Council for providing the funds which enabled this work to be carried out. R.M.G.W. was further supported by a grant from the Auckland University Research. References Birchard G. F., Black C. P., Schuett G. W. and Black V. (1984) Foetal-maternal blood respiratory properties of an ovoviviparous snake the cottonmouth, Agkistrodon piscivorous. J. Exp. Biol. 108, Chien S., Usami S., Dellenmack D. J. and Bryant C. A. (1971) Comparative hemorheologyhematological implications of species differences in blood viscosity. Biorheology 8, Crowell J. W. and Smith E. E. (1967) Determinant of the optimal hematocrit. J. Appl. Physiol. 22, Dacie J. V. and Lewis S. N. (1984) Practical Haematology 5th edition, 453 pp. Churchill Livingstone, Edinburgh. Dawson W. R. and Poulson T. L. (1962) Oxygen capacity of lizard bloods. Am. Midl. Nat. 68, Fletcher G. L. and Haedrich R. T. (1987) Rheological properties of rainbow trout blood. Can. J. Zoo/. 65, Gatten R. E. (1985) The uses of anaerobiosis by amphibians and reptiles. Amer. Zoo/. 25, Glass M. L. and Johansen K. (1979) Periodic breathing in the crocodile, Crocodylus niloticus: consequences for the gas exchange ratio and control of breathing. J. Exp. Zool. 208,
6 Glass M. L. and Wood S. C. (1983) Gas exchange and control of breathing in reptiles. Physiol. Rev. 63, Grigg G. C. (1978) Metabolic rate, Qio and respiratory quotient (RQ) in Crocodylus porosus, and some generalizations about low RQ in reptiles. Physiol. Zool. 51, Grigg G. (1989) The heart and patterns of cardiac outflow in Crocodilia. Proc. Aust. Physiol. Pharmacol. Soc. 20, Grigg G. C. and Cairncross M. (1980) Respiratory proper-ties of the blood of Crocodylus porosus. Respir. Physiol. 41, Grigg G. C. and Gruca M. (1979) Possible adaptive significance of low red cell organic phosphates in crocodiles. J. Exp. Zool. 209, Grigg G. C. and Johansen K. (1987) Cardiovascular dynamics in Crocodylus porosus breathing air and during voluntary aerobic dives. J. Comp. Physiol. B 157, Grigg G. C., Farwell W. D., Kinney J. L., Harlow P., Taplin, L. E., Johansen, Kjell and Johansen, Kjetil (1985) Diving and amphibious behaviour in a free-living Crocodylus porosus. Aust. Zoo/. 21, Gruca M. and Grigg G. C. (1980) Methemoglobin reduction in crocodile blood: are high levels of met Hb typical of healthy reptiles? J. Exp. Zool. 213, Hedrick M. S., Duffield D. A. and Cornell L. H. (1986) Blood viscosity and optimal hematocrit in a deep-diving mammal, the northern elephant seal (Mirounga angustirostris). Can. J. Zoo/. 64, Hillman S. S., Withers P. C., Hedrick M. S. and Kimmel P. B. (1985) The effects of erythrocythemia on blood viscosity, maximal systemic oxygen transport capacity and maximal rates of oxygen consumption in an amphibian. J. Comp. Physiol, B 155, Jelkmann W. and Bauer C. (1980) Oxygen binding proper-ties of caiman blood in the presence and absence of carbon dioxide. Comp. Biochem. Physiol. 65A, Kooyman G. L., Castellini M. A. and Davis R. W. (1981) Physiology of diving in marine mammals. Ann. Rev. Physiol. 43, Lewis, L. Y. and Gatten R. E. (1985) Aerobic metabolism of American alligators. Alligator mississippiensis, under standard conditions and showing voluntary activity. Comp. Biochem. Physiol. 80A, Lutz P. L. and Bentley T. B. (1985) Respiratory physiology of diving in the sea turtle. Copeia 1985, Maginniss L. A., Tapper S. S. and Miller L. S. (1983) Effect of chronic cold and submergence on blood oxygen trans-port in the turtle, Chrysemys picta. Respir. Physiol. 53, Pough H. (1980) Blood oxygen transport and delivery in reptiles. Amer. Zoo/. 20, Rand P. W., Lacombe E., Hunt H. E. and Austin W. H. (1964) Viscosity of normal human blood under normothermic and hypothermic conditions. J. Appl. Physiol. 19, Seymour R. S., Bennett A. F. and Bradford D. F. (1985) Blood gas tension and acid-base regulation in the salt-water crocodile, Crocodylus porosus, at rest and after exhaustive exercise. J. Exp. Biol. 118, Seymour R. S., Dobson G. P. and Baldwin J. (1981) Respiratory and cardiovascular physiology of the aquatic snake, Acrochordus arafurae. J. Comp. Physiol. 144, Snyder G. K. (1983) Respiratory adaptations in diving mammals. Resp. Physiol. 54, Weathers W. W. (1976) Influence of temperature on the optimal hematocrit of the bullfrog (Rana catesbeiana). J. Comp. Physiol. 105, Wells R. M. G. and Forster M. E. (1989) Dependence of blood viscosity on haematocrit and shear rate in a primitive vertebrate. J. Exp. Biol. 145, Wickham L. L., Elsner R., White F. C. and Cornell L. H. (1989) Blood viscosity in phocid seals: possible adaptations to diving. J. Comp. Physiol. B 159, Wood S. C. and Johansen K. (1974) Respiratory adaptations to diving in the Nile monitor lizard, Varanus niloticus. J. Comp. Physiol. 89,
Respiration Physiology (1980) RESPIRATORY PROPERTIES OF THE BLOOD OF CROCODYLUS POROSUS GORDON C. GR1GG and MICHAEL CAIRNCROSS
Respiration Physiology (1980) 41. 367-380 RESPIRATORY PROPERTIES OF THE BLOOD OF CROCODYLUS POROSUS GORDON C. GR1GG and MICHAEL CAIRNCROSS Abstract. The blood of Crocodylus porosus has a high oxygen capacity
More informationJeff Baier MS DVM Birds of Prey Foundation Broomfield, CO
Jeff Baier MS DVM Birds of Prey Foundation Broomfield, CO drjeffbaier@gmail.com Squamates Chelonians Snakes Lizards Varanids Monitor Lizards Crocodilians Reptilian adaptations Anaerobic glycolysis Low
More informationTopic 13: Energetics & Performance. How are gas exchange, circulation & metabolism inter-related?
Topic 3: Energetics & Performance How are gas exchange, circulation & metabolism interrelated? How is it done in air and water? What organs are involved in each case? How does ventilation differ among
More informationMechanism of a Crocodile s Circulatory System
Mechanism of a Crocodile s Circulatory System Figure 1. A crocodile diving at Botswana (Nachoum, A. 2017) Ever wonder in one of those animal documentaries we watch in television, wherein a crocodile glides
More informationFACULTATIVE AESTIVATION IN A TROPICAL FRESHWATER TURTLE CHELODINA RUGOSA
FACULTATIVE AESTIVATION IN A TROPICAL FRESHWATER TURTLE CHELODINA RUGOSA G. C. GRIGG, * K. JOHANSEN, P. HARLOW, * L. A. BEARD* and L. E. TAPLIN *Zoology A.08, The University of Sydney, NSW 2006, Australia.
More informationBREATHING WHICH IS NOT RESPIRATION
BREATHING WHICH IS NOT RESPIRATION Breathing vs. Respiration All animals respire. A lot of people think respiration means breathing- this is not true! Breathing is the physical process of inhaling oxygen
More informationComparative Physiology 2007 Second Midterm Exam. 1) 8 pts. 2) 14 pts. 3) 12 pts. 4) 17 pts. 5) 10 pts. 6) 8 pts. 7) 12 pts. 8) 10 pts. 9) 9 pts.
Name: Comparative Physiology 2007 Second Midterm Exam 1) 8 pts 2) 14 pts 3) 12 pts 4) 17 pts 5) 10 pts 6) 8 pts 7) 12 pts 8) 10 pts 9) 9 pts Total 1. Cells I and II, shown below, are found in the gills
More informationEat and run: prioritization of oxygen delivery during elevated metabolic states
Respiratory Physiology & Neurobiology 144 (2004) 215 224 Eat and run: prioritization of oxygen delivery during elevated metabolic states James W. Hicks, Albert F. Bennett Department of Ecology and Evolutionary
More informationVertebrates. skull ribs vertebral column
Vertebrates skull ribs vertebral column endoskeleton in cells working together tissues tissues working together organs working together organs systems Blood carries oxygen to the cells carries nutrients
More informationSec KEY CONCEPT Reptiles, birds, and mammals are amniotes.
Thu 4/27 Learning Target Class Activities *attached below (scroll down)* Website: my.hrw.com Username: bio678 Password:a4s5s Activities Students will describe the evolutionary significance of amniotic
More informationReptilian Requirements Created by the North Carolina Aquarium at Fort Fisher Education Section
Essential Question: North Carolina Aquariums Education Section Reptilian Requirements Created by the North Carolina Aquarium at Fort Fisher Education Section What physical and behavioral adaptations do
More informationBiology. Slide 1of 50. End Show. Copyright Pearson Prentice Hall
Biology 1of 50 2of 50 Phylogeny of Chordates Nonvertebrate chordates Jawless fishes Sharks & their relatives Bony fishes Reptiles Amphibians Birds Mammals Invertebrate ancestor 3of 50 A vertebrate dry,
More informationFrom Slime to Scales: Evolution of Reptiles. Review: Disadvantages of Being an Amphibian
From Slime to Scales: Evolution of Reptiles Review: Disadvantages of Being an Amphibian Gelatinous eggs of amphibians cannot survive out of water, so amphibians are limited in terms of the environments
More informationQuestion Set 1: Animal EVOLUTIONARY BIODIVERSITY
Biology 162 LAB EXAM 2, AM Version Thursday 24 April 2003 page 1 Question Set 1: Animal EVOLUTIONARY BIODIVERSITY (a). We have mentioned several times in class that the concepts of Developed and Evolved
More informationCharacteristics of Tetrapods
Marine Tetrapods Characteristics of Tetrapods Tetrapod = four-footed Reptiles, Birds, & Mammals No marine species of amphibian Air-breathing lungs Class Reptilia Saltwater Crocodiles, Sea turtles, sea
More informationBiology Slide 1 of 50
Biology 1 of 50 2 of 50 What Is a Reptile? What are the characteristics of reptiles? 3 of 50 What Is a Reptile? What Is a Reptile? A reptile is a vertebrate that has dry, scaly skin, lungs, and terrestrial
More informationRELATIONSHIP BETWEEN HAEMOGLOBIN O 2 AFFINITY AND THE VENTILATORY RESPONSE TO HYPOXIA IN THE RHEA AND PHEASANT
J. exp. Biol. 102, 347352, 1983 347 ^Printed in Great Britain Company of Biologists Limited 1983 RELATIONSHIP BETWEEN HAEMOGLOBIN O 2 AFFINITY AND THE VENTILATORY RESPONSE TO HYPOXIA IN THE RHEA AND PHEASANT
More informationEFFECTS OF TEMPERATURE ON GAS EXCHANGE AND ACID-BASE BALANCE IN THE SEA TURTLE CARETTA CARETTA AT REST AND DURING ROUTINE ACTIVITY
/. exp. Biol. 144, 155-169 (1989) 155 Printed in Great Britain The Company of Biologists Limited 1989 EFFECTS OF TEMPERATURE ON GAS EXCHANGE AND ACID-BASE BALANCE IN THE SEA TURTLE CARETTA CARETTA AT REST
More informationVERTEBRATE READING. Fishes
VERTEBRATE READING Fishes The first vertebrates to become a widespread, predominant life form on earth were fishes. Prior to this, only invertebrates, such as mollusks, worms and squid-like animals, would
More informationKINGDOM ANIMALIA Phylum Chordata Subphylum Vertebrata Class Reptilia
KINGDOM ANIMALIA Phylum Chordata Subphylum Vertebrata Class Reptilia Vertebrate Classes Reptiles are the evolutionary base for the rest of the tetrapods. Early divergence of mammals from reptilian ancestor.
More informationOXYGEN AND CARBON DIOXIDE TRANSPORT CHARACTERISTICS OF THE BLOOD OF THE NILE MONITOR LIZARD (VARANUS NILOTICUS)
J. exp. Biol. 130, 27-38 (1987) 27 Printed in Great Britain The Company of Biologists Limited 1987 OXYGEN AND CARBON DOXDE TRANSPORT CHARACTERSTCS OF THE BLOOD OF THE NLE MONTOR LZARD (VARANUS NLOTCUS)
More informationGAS EXCHANGE AND VENTILATORY RESPONSES TO HYPOXIA AND HYPERCAPNIA IN AMPHISBAENA ALBA (REPTILIA: AMPHISBAENIA)
J. exp. Biol. 127, 159-172 (1987) \ 59 Printed in Great Britain The Company of Biologists Limited 1987 GAS EXCHANGE AND VENTILATORY RESPONSES TO HYPOXIA AND HYPERCAPNIA IN AMPHISBAENA ALBA (REPTILIA: AMPHISBAENIA)
More informationConservation (last three 3 lecture periods, mostly as a led discussion). We can't cover everything, but that should serve as a rough outline.
Comments on the rest of the semester: Subjects to be discussed: Temperature relationships. Echolocation. Conservation (last three 3 lecture periods, mostly as a led discussion). Possibly (in order of importance):
More informationPASSIVE BODY MOVEMENT AND GAS EXCHANGE IN THE FRILLED LIZARD (CHLAMYDOSAURUS KINGII) AND GOANNA (VARANUS GOULDII)
The Journal of Experimental Biology, 7 (998) Printed in Great Britain The Company of Biologists Limited 998 JEB7 7 PASSIVE BODY MOVEMENT AND GAS EXCHANGE IN THE FRILLED LIZARD (CHLAMYDOSAURUS KINGII) AND
More informationThe cardiovascular responses of the freshwater turtle Trachemys scripta to warming and cooling
The Journal of Experimental Biology 27, 1471-1478 Published by The Company of Biologists 24 doi:1.1242/jeb.912 1471 The cardiovascular responses of the freshwater turtle Trachemys scripta to warming and
More informationName Date When you put food away in the kitchen, you sort the food into groups. You put foods that are alike in certain ways into the same
1 Name Date When you put food away in the kitchen, you sort the food into groups. You put foods that are alike in certain ways into the same group. Scientists do the same thing with animals, plants and
More informationInvestigating Fish Respiration
CHAPTER 31 Fishes and Amphibians Section 31-1 SKILL ACTIVITY Interpreting graphs Investigating Fish Respiration It is well known that a fish dies from lack of oxygen when taken out of water. However, water
More informationT. 6. THE VERTEBRATES
T. 6. THE VERTEBRATES 1.- Relate the following concepts to their definition. Later, relate each concept to one of the pictures you are going to see. 1.- FIN a.- mammals with their babies 2.- GILLS b.-
More informationPhylogeny of Animalia (overview)
The Diversity of Animals 2 Chapter 23 Phylogeny of Animalia (overview) Key features of Chordates Phylum Chordata (the Chordates) includes both invertebrates and vertebrates that share (at some point in
More informationDiversity of Animals
Classifying Animals Diversity of Animals Animals can be classified and grouped based on similarities in their characteristics. Animals make up one of the major biological groups of classification. All
More informationREPTILES. Scientific Classification of Reptiles To creep. Kingdom: Animalia Phylum: Chordata Subphylum: Vertebrata Class: Reptilia
Scientific Classification of Reptiles To creep Kingdom: Animalia Phylum: Chordata Subphylum: Vertebrata Class: Reptilia REPTILES tetrapods - 4 legs adapted for land, hip/girdle Amniotes - animals whose
More informationAnimal Diversity wrap-up Lecture 9 Winter 2014
Animal Diversity wrap-up Lecture 9 Winter 2014 1 Animal phylogeny based on morphology & development Fig. 32.10 2 Animal phylogeny based on molecular data Fig. 32.11 New Clades 3 Lophotrochozoa Lophophore:
More informationVertebrate and Invertebrate Animals
Vertebrate and Invertebrate Animals Compare the characteristic structures of invertebrate animals (including sponges, segmented worms, echinoderms, mollusks, and arthropods) and vertebrate animals (fish,
More informationACTIVITY METABOLISM
Ann. Rev. Physiol. 1978. 400:447-69 Copyright O 1978 by Annual Reviews Inn. AN righrs reserved ACTIVITY METABOLISM +1 198 OF THE LOWER VERTEBRATES Albert l? Bennett School of Biological Sciences, University
More information5 pt. 10 pt. 15 pt. 20 pt. 25 pt
Final Jeopardy Characteristics of Vertebrates Characteristics of Fish Amphibians Reptiles Chapter 16 Vocabulary 5 pt 5 pt 5 pt 5 pt 5 pt 10 pt 10 pt 10 pt 10 pt 10 pt 15 pt 15 pt 15 pt 15 pt 15 pt 20 pt
More informationClass Reptilia Testudines Squamata Crocodilia Sphenodontia
Class Reptilia Testudines (around 300 species Tortoises and Turtles) Squamata (around 7,900 species Snakes, Lizards and amphisbaenids) Crocodilia (around 23 species Alligators, Crocodiles, Caimans and
More informationReturn to the sea: Marine birds, reptiles and pinnipeds
Figure 34.14 The origin of tetrapods Return to the sea: Marine birds, reptiles and pinnipeds Phylum Chordata Free swimmers Nekton Now we move to reptiles (Class Reptilia) and birds (Class Aves), then on
More informationSummary. Introduction
Grigg GC, LE Taplin, P Harlow and J Wright 1980 Survival and growth of hatchling Crocodylus porosus in salt water without access to fresh drinking water. Oecologia 47:264-6. Survival and Growth of Hatchling
More informationWhat does it mean to be a tetrapod? What three things were needed to survive on land? What does it mean to be oviparous?
Marine Mammals, Reptiles, Amphibians and Birds What does it mean to be a tetrapod? What three things were needed to survive on land? Which two of these problems did amphibians NOT figure out? What does
More informationA REAPPRAISAL OF THE AQUATIC SPECIALIZATIONS OF THE GALAPAGOS MARINE IGUANA (AMBLYRHYNCHUS CRISTATUS)
A REAPPRAISAL OF THE AQUATIC SPECIALIZATIONS OF THE GALAPAGOS MARINE IGUANA (AMBLYRHYNCHUS CRISTATUS) Wn.LIAM R. DAWSON, GEORGE A. BARTHOLOMEW, AND ALBERT F. BENNETT Division of Biological Sciences, The
More informationVertebrates. Vertebrate Characteristics. 444 Chapter 14
4 Vertebrates Key Concept All vertebrates have a backbone, which supports other specialized body structures and functions. What You Will Learn Vertebrates have an endoskeleton that provides support and
More informationOsmoregulation Chapter 26 & 27
31 st Lecture Fri 03 April 2009 Vertebrate Physiology ECOL 437 (MCB/VetSci 437) Univ. of Arizona, spring 2009 Kevin Bonine & Kevin Oh Housekeeping, Wed 01 April 2009 Readings Today, Mon 30 Mar: Ch 26 (Ionic
More informationOsmoregulation. 31 st Lecture Fri 03 April Chapter 26 & 27. Research Proposal Meetings 1
31 st Lecture Fri 03 April 2009 Vertebrate Physiology ECOL 437 (MCB/VetSci 437) Univ. of Arizona, spring 2009 Kevin Bonine & Kevin Oh Osmoregulation Chapter 26 & 27 Research Proposal Meetings 1 Housekeeping,
More informationMASS-DEPENDENCE OF ANAEROBIC METABOLISM AND ACID-BASE DISTURBANCE DURING ACTIVITY IN THE SALT-WATER CROCODILE, CROCODYLUS POROSUS
Jf. exp. Biol. 118, 161-171 (1985) 161 Printed in Great Britain The Company of Biologists Limited 1985 MASS-DEPENDENCE OF ANAEROBIC METABOLISM AND ACID-BASE DISTURBANCE DURING ACTIVITY IN THE SALT-WATER
More informationThe Saltwater Crocodile Crocodylus porosus (Schneider, 1801)
The Saltwater Crocodile Crocodylus porosus (Schneider, 1801) Taxonomy Kingdom Animalia Phylum - Chordata Class - Sauropsida Order - Crocodilia Family Crocodylidae Subfamily - Crocodylinae Genus - Crocodylus
More informationBROOD REDUCTION IN THE CURVE-BILLED THRASHER By ROBERTE.RICKLEFS
Nov., 1965 505 BROOD REDUCTION IN THE CURVE-BILLED THRASHER By ROBERTE.RICKLEFS Lack ( 1954; 40-41) has pointed out that in species of birds which have asynchronous hatching, brood size may be adjusted
More informationCharacteristics of a Reptile. Vertebrate animals Lungs Scaly skin Amniotic egg
Reptiles Characteristics of a Reptile Vertebrate animals Lungs Scaly skin Amniotic egg Characteristics of Reptiles Adaptations to life on land More efficient lungs and a better circulator system were develope
More informationLactic Acid Buffering by Bone and Shell in Anoxic Softshell and Painted Turtles
290 Lactic Acid Buffering by Bone and in Anoxic Softshell and Painted Turtles D. C. Jackson 1,* A. L. Ramsey 1 J. M. Paulson 1 C. E. Crocker 1,2 G. R. Ultsch 2 1 Department of Molecular Pharmacology, Physiology,
More informationIs Aquatic Life Correlated with an Increased Hematocrit in Snakes?
Is Aquatic Life Correlated with an Increased Hematocrit in Snakes? François Brischoux 1 *, Gabriel E. A. Gartner 2, Theodore Garland Jr. 2, Xavier Bonnet 1 1 Centre d Etudes Biologiques de Chizé CNRS,
More informationWhat is the body structure of a sponge? Do they have specialized cells? Describe the process of reproduction in sponges.
11.2 Sponges and Cnidarians What are the main characteristics of Sponges? Where are sponges found? What is the body structure of a sponge? Do they have specialized cells? Do sponges have separate sexes?
More informationCurrent Status of Amphibian Populations. Amphibian biology - characteristics making
Global Amphibian Declines: What Have We Done? Mike Tyler Steve Holmer Nikki Maxwell University of Tennessee Knoxville Department of Forestry, Wildlife and Fisheries Graduate Student Seminar 15 October
More informationACID-BASE STATUS OF BLOOD OF V ARANUS GRISEUS AND UROMASTYX AEGYPTIUS
Qatar Univ. Sci. Bull. (1984) 4: 159-170 ACID-BASE STATUS OF BLOOD OF V ARANUS GRISEUS AND UROMASTYX AEGYPTIUS By SAID M. EISSA* and WAFAA S. HASHEESH Department of Zoology, Faculty of Science, Cairo University,
More information4 Many species of mammals, birds, reptiles, amphibians and fish 940L. Source 1 Habitats
Source 1 Habitats 1 American Alligators can be found in fresh water environments like rivers, lakes, ponds, swamps and marshes. They also like to live in areas that are brackish, which means the water
More informationOXYGEN POISONING IN COLD BLOODED ANIMALS, By JAMES M. FAULKNER, M.D., AND CARL A. L. BINGER, M.D. (Received for publication, January 3, 1927.
Published Online: 1 May, 1927 Supp Info: http://doi.org/10.1084/jem.45.5.865 Downloaded from jem.rupress.org on September 21, 2018 OXYGEN POISONING IN COLD BLOODED ANIMALS, By JAMES M. FAULKNER, M.D.,
More informationSome important information about the fetus and the newborn puppy
Some important information about the fetus and the newborn puppy Dr. Harmon Rogers Veterinary Teaching Hospital Washington State University Here are a few interesting medical details about fetuses and
More information2/11/2015. Body mass and total Glomerular area. Body mass and medullary thickness. Insect Nephridial Structure. Salt Gland Structure
Body mass and medullary thickness Thicker medulla in mammals from dry climate Negative allometry why? Body mass and total Glomerular area Glomerular area is a measure of total ultrafiltration rate Slope
More informationAustralian and New Zealand College of Veterinary Scientists. Membership Examination. Veterinary Anaesthesia and Critical Care Paper 1
Australian and New Zealand College of Veterinary Scientists Membership Examination June 2015 Veterinary Anaesthesia and Critical Care Paper 1 Perusal time: Fifteen (15) minutes Time allowed: Two (2) hours
More informationEssential Question: What are the characteristics of invertebrate animals? What are the characteristics of vertebrate animals?
Essential Question: What are the characteristics of invertebrate animals? What are the characteristics of vertebrate animals? Key Concept: The animal kingdom is divided up into 35 phyla. These phyla can
More informationAustralian and New Zealand College of Veterinary Scientists. Fellowship Examination. Veterinary Anaesthesia and Critical Care Paper 1
Australian and New Zealand College of Veterinary Scientists Fellowship Examination June 2016 Veterinary Anaesthesia and Critical Care Paper 1 Perusal time: Twenty (20) minutes Time allowed: Three (3) hours
More informationVertebrate Structure and Function
Vertebrate Structure and Function Part 1 - Comparing Structure and Function Classification of Vertebrates a. Phylum: Chordata Common Characteristics: Notochord, pharyngeal gill slits, hollow dorsal nerve
More informationAPNOEA IN AMPHIBIANS AND REPTILES
J. exp. Biol. (1982), ioo, 245-273 245 With 12 figures (in Great Britain APNOEA IN AMPHIBIANS AND REPTILES BY G. SHELTON AND R. G. BOUTILIER School of Biological Sciences, University of East Anglia, Norwich,
More informationField Lesson: Reptiles and Amphibians
Field Lesson: Reptiles and Amphibians State Core Standards 5.2 Interaction and Change: Force, energy, matter, and organisms interact within living and non-living systems Content Standards 5.2L.1 Explain
More informationAnimal Form and Function. Amphibians. United by several distinguishing apomorphies within the Vertebrata
Animal Form and Function Kight Amphibians Class Amphibia (amphibia = living a double life) United by several distinguishing apomorphies within the Vertebrata 1. Skin Thought Question: For whom are integumentary
More information13. Swim bladder function: A. What happens to the density of a fish if the volume of its swim bladder increases?
Ch 11 Review - Use this worksheet as practice and as an addition to your Chapter 11 Study Guide. Test will only be over Ch 11.1-11.4. (Ch 11.5 Fossil and Paleontology section will not be on your test)
More informationThe critical importance of incubation temperature
The critical importance of incubation temperature Nick A. French AVIAN BIOLOGY RESEARCH 2 (1/2), 2009 55 59 Aviagen Turkeys Ltd, Chowley Five, Chowley Oak Business Park, Tattenhall, Cheshire, CH3 9GA,
More informationBehavioral and Physiological Thermoregulation of Crocodilians
AMER. ZOOL..19:239-247 (1979). Behavioral and Physiological Thermoregulation of Crocodilians E. NORBERT SMITH Northeastern Oklahoma State University, Tahlequah, Oklahoma 74464 SYNOPSIS. Crocodilians, like
More informationVertebrates. Vertebrates are animals that have a backbone and an endoskeleton.
Vertebrates Vertebrates are animals that have a backbone and an endoskeleton. The backbone replaces the notochord and contains bones called vertebrae. An endoskeleton is an internal skeleton that protects
More informationIMPORT HEALTH STANDARD FOR ZOO CROCODILIA HATCHING EGGS FROM AUSTRALIA
IMPORT HEALTH STANDARD FOR ZOO CROCODILIA HATCHING EGGS FROM AUSTRALIA Issued pursuant to Section 22 of the Biosecurity Act 1993 Dated: 21 April 2008 USER GUIDE The information in MAFBNZ animal and animal
More informationOxygen transport in varanid lizards during exercise. Timothy John Schultz
Oxygen transport in varanid lizards during exercise Timothy John Schultz B.A. (Adelaide), B. Sc. (Hons) (Adelaide) A thesis submitted to satisfy the requirements for the a~ard of the degree of Doctor ofphilosophy
More informationFishes, Amphibians, Reptiles
Fishes, Amphibians, Reptiles Section 1: What is a Vertebrate? Characteristics of CHORDATES Most are Vertebrates (have a spinal cord) Some point in life cycle all chordates have: Notochord Nerve cord that
More informationThe Friends of Nachusa Grasslands 2016 Scientific Research Project Grant Report Due June 30, 2017
The Friends of Nachusa Grasslands 2016 Scientific Research Project Grant Report Due June 30, 2017 Name: Laura Adamovicz Address: 2001 S Lincoln Ave, Urbana, IL 61802 Phone: 217-333-8056 2016 grant amount:
More informationName Class Date. After you read this section, you should be able to answer these questions:
CHAPTER 14 4 Vertebrates SECTION Introduction to Animals BEFORE YOU READ After you read this section, you should be able to answer these questions: How are vertebrates different from invertebrates? How
More information8/19/2013. Topic 12: Water & Temperature. Why are water and temperature important? Why are water and temperature important?
Topic 2: Water & Temperature Why are water and temperature important? Why are water and temperature important for herps? What are adaptations for gaining water? What are adaptations for limiting loss of
More information#8964 Standards-Based Science Investigations 2 Teacher Created Resources, Inc.
Introduction...4 Locating Simple Science Materials...5 Standards Correlation....7 Thinking About Inquiry Investigations...9 Inquiry Assessment Rubric...12 Student Inquiry Worksheets...13 Sample Inquiry
More informationFeline blood transfusions: preliminary considerations
Vet Times The website for the veterinary profession https://www.vettimes.co.uk Feline blood transfusions: preliminary considerations Author : Andrea Harvey Categories : RVNs Date : September 1, 2011 ABSTRACT
More information*Using the 2018 List. Use the image below to answer question 6.
Herpetology Test 1. Hearts in all herps other than consists of atria and one ventricle somewhat divided by a septum. (2 pts) a. snakes; two b. crocodiles; two c. turtles; three d. frogs; four 2. The food
More informationExceptions: Somebody liked snakes. Some people disliked dogs, geese, sharks
Unit 1: ANIMALS Exceptions: Somebody liked snakes Some people disliked dogs, geese, sharks Both animals are fascinating & worthy of our interest ANIMAL NAMES Taxonomy is a branch of biology that categorizes
More informationA. Body Temperature Control Form and Function in Mammals
Taxonomy Chapter 22 Kingdom Animalia Phylum Chordata Class Mammalia Mammals Characteristics Evolution of Mammals Have hair and First appear in the mammary glands Breathe air, 4chambered heart, endotherms
More informationThe effect of body temperature on the locomotory energetics of lizards
J Comp Physiol B (1984) 155: 21-27 Journal of @ Springer-Verlag 1984 The effect of body temperature on the locomotory energetics of lizards Albert F. Bennett and Henry B. John-Alder School of Biological
More informationRookery on the east coast of Penins. Author(s) ABDULLAH, SYED; ISMAIL, MAZLAN. Proceedings of the International Sy
Temperature dependent sex determina Titleperformance of green turtle (Chelon Rookery on the east coast of Penins Author(s) ABDULLAH, SYED; ISMAIL, MAZLAN Proceedings of the International Sy Citation SEASTAR2000
More informationReptilian Physiology
Reptilian Physiology Physiology, part deux The study of chemical and physical processes in the organism Aspects of the physiology can be informative for understanding organisms in their environment Thermoregulation
More informationLingual Salt Glands in Crocodylus acutus and C. johnstoni and their absence from Alligator mississipiensis and Caiman crocodilus
Lingual Salt Glands in Crocodylus acutus and C. johnstoni and their absence from Alligator mississipiensis and Caiman crocodilus Laurence E. Taplin 1, Gordon C. Grigg 1, Peter Harlow 1, Tamir M. Ellis
More informationPhilippines Crocodile (Crocodylus mindorensis ) - the effects of temperature on sex determination.
% of females Introduction: The Philippines Crocodile is a critically endangered species found in a small range of islands in the Philippines. The species is under threat from habitat destruction and practices
More informationRed Eared Slider Secrets. Although Most Red-Eared Sliders Can Live Up to Years, Most WILL NOT Survive Two Years!
Although Most Red-Eared Sliders Can Live Up to 45-60 Years, Most WILL NOT Survive Two Years! Chris Johnson 2014 2 Red Eared Slider Secrets Although Most Red-Eared Sliders Can Live Up to 45-60 Years, Most
More informationToday s Class. Go over viewfinder lab A closer look at the Animal Kingdom Taxonomy Worksheet
Kingdom Animalia Today s Class Go over viewfinder lab A closer look at the Animal Kingdom Taxonomy Worksheet Viewfinder Kingdom: Animalia Cells: multicellular Food: eats other organisms Movement: moves
More informationSome Facts about... Amphibians
Amphibians Amphibians are cold-blooded vertebrates that live part of their lives in water and part on land. Amphibians eggs are laid in water and they are born there. They begin their lives with gills
More informationRecovery from an activity-induced metabolic acidosis in the American alligator, Alligator mississippiensis
Comparative Biochemistry and Physiology, Part A 143 (6) 368 374 www.elsevier.com/locate/cbpa Recovery from an activity-induced metabolic acidosis in the American alligator, Alligator mississippiensis L.K.
More informationReptiles Amphibians ( am-fib-ee-anz ) Fish Birds Mammals
Chapter 11: Page 94 In the last chapter, you learned how plants go through a natural cycle of life. It is not just plants that go through a life cycle! Animals do too! Remember the definition of species?
More informationGrade Level: 1-2. Next Generation Sunshine State Standards SC.1.L.14.1; SC.1.L.17.1; SC.1.N.1.1 SC.2.L.17.1; SC.2.L.17.2; SC.2.N.1.
Grade Level: 1-2 Next Generation Sunshine State Standards SC.1.L.14.1; SC.1.L.17.1; SC.1.N.1.1 SC.2.L.17.1; SC.2.L.17.2; SC.2.N.1.1 Program Overview Reptiles Rock! Meet live reptiles up close and investigate
More informationMost amphibians begin life as aquatic organisms and then live on land as adults.
Section 3: Most amphibians begin life as aquatic organisms and then live on land as adults. K What I Know W What I Want to Find Out L What I Learned Essential Questions What were the kinds of adaptations
More informationGet the other MEGA courses!
www.thesimplehomeschool.com Simple Schooling BUGS MEGA course is ten weeks of all about bugs! This course grabs your student s attention and never lets go! Grades K-3 Get the other MEGA courses! Simple
More informationBig Picture Biology Student Workbook
Big Picture Biology Student Workbook VCE Biology Unit 1 & 2 What do we need to know about an endangered species in order to save it? Big Picture Biology provides a unique insight into how the VCE biology
More informationFederal Department of Economic Affairs FDEA Federal Veterinary Office FVO. Need for global animal welfare. standards for reptiles
Federal Department of Economic Affairs FDEA Federal Veterinary Office FVO Need for global animal welfare standards for reptiles M. Lörtscher, Head CITES MA Switzerland, Import & keeping of exotic animals
More informationDIFFERENT BREEDS DEMAND DIFFERENT INCUBATION MEASURES
CONCERNING POULTRY One can be puzzled by noticing that, from the same batch, in the same incubator, some of the chicks hatch normally, while others die before breaking the shell. Reading the following
More informationAmphibians. Land and Water Dwellers
Amphibians Land and Water Dwellers Amphibians Most amphibians do not live completely in the water or completely on land and most must return to water to reproduce http://potch74.files.wordpress.com/2007/09/amphibians.jpg
More informationLiving Dinosaurs (3-5) Animal Demonstrations
Living Dinosaurs (3-5) Animal Demonstrations At a glance Students visiting the zoo will be introduced to live animals and understand their connection to a common ancestor, dinosaurs. Time requirement One
More informationTaxonomy. Chapter 20. Evolutionary Development Diagram. I. Evolution 2/24/11. Kingdom - Animalia Phylum - Chordata Class Reptilia.
Taxonomy Chapter 20 Reptiles Kingdom - Animalia Phylum - Chordata Class Reptilia Order Testudines - turtles Order Crocodylia - crocodiles, alligators Order Sphenodontida - tuataras Order Squamata - snakes
More informationGrade Level: 3-5. Next Generation Sunshine State Standards SC.3.L.15.1 SC.4.L.16.2; SC.4.L.17.4 SC.5.L.15.1; SC.5.L.17.1
Grade Level: 3-5 Next Generation Sunshine State Standards SC.3.L.15.1 SC.4.L.16.2; SC.4.L.17.4 SC.5.L.15.1; SC.5.L.17.1 Program Overview Discover the realm of reptiles, amazing creatures adapted to land
More information1. Examine the specimens of sponges on the lab table. Which of these are true sponges? Explain your answers.
Station #1 - Porifera 1. Examine the specimens of sponges on the lab table. Which of these are true sponges? Explain your answers. 2. Sponges are said to have an internal special skeleton. Examine the
More informationOxygen. Carbon Dioxide. Carbon Dioxide. Oxygen. Aquatic Plants. Fish
Aquaponics System: A fish tank is an example of an aquaponics ecosystem. In an aquaponics ecosystem, a sustainable food production cycle is created through the interaction of the animals and plants within
More information