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Haematology of Amphibians and Reptiles: a Review

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Haematology of Amphibians and Reptiles: a Review NORTH-WESTERN JOURNAL OF ZOOLOGY 10 (1): 190-209 ©NwjZ, Oradea, Romania, 2014 Article No.: 143501 http://biozoojournals.ro/nwjz/index.html Haematology of amphibians and reptiles: a review Hüseyin ARIKAN and Kerim ÇİÇEK* Ege University, Faculty of Science, Biology Department, Zoology Section, TR-35100, Bornova-Izmir, Turkey. *Corresponding author, K. Çiçek, E-mail’s: [email protected], [email protected] Received: 22. January 2013 / Accepted: 19. October 2013 / Available online: 28. January 2014 / Printed: June 2014 Abstract. In this review, function and composition of blood, haemopoiesis, morphology, number and size of blood cells (erythrocytes, leukocytes and thrombocytes), and intra/extracellular blood parasites of amphibians and reptiles are revised by utilizing the literature knowledge. Key words: Amphibia, Reptilia, blood cells, plasma, haemopoiesis, blood parasites. Introduction from mesonephros type kidney. Until the comple- tion of metamorphosis, spleen is also taking on the In old Greek, haemat means blood and /o/-logy task of creating new blood cells (Jordan 1933). means investigation or examination. Thus, haema- Most of the erythrocytes, thrombocytes and leuko- tology refers to the examination of blood (Seiverd cytes are formed in the bone marrow in adults. 1972). Multicellular animal organisms need the The most widespread sources of the red bone mar- circulatory system to transport food particles and row are ribs, sternum and spine (Seiverd 1972). respiratory gases to the cells in tissues and to carry the wastes from cells to the necessary organs. Hematopoiesis in Amphibians Blood is the circulatory fluid in vertebrates. Keep- Haemopoiesis in amphibians has been studied by ing both the components and the amount of blood many researchers (e.g. Jordan 1938, Jordan & at a constant level is essential for the continuity of Speidel 1923, Maniatis & Ingram 1971, Hadji- life. Azimi et al. 1987, Allander & Fry 2008). The de- velopment of hematopoiesis in amphibians dis- plays some similarities to those of birds and 1. Characteristics of Vertebrates Blood mammals (Allander & Fry 2008). There are great differences in erythropoiesis between juvenile and Composition of Blood adult amphibians. Erythropoiesis takes place in Blood corpuscles, or blood cells (erythrocytes, leuko- the liver and kidney in juvenile amphibians, while cytes and thrombocytes), and the liquid intercellular it takes place in the bone marrow as well as in the substance are a type of specialized connective tis- spleen and the liver in adult aquatic and terrestrial sue of mesenchymal origin originating from the frogs. Thyroxine application affects erythropoiesis blood plasma. Plasma consists of water, food sub- in adults. In amphibians, ventral blood islets are stances, wastes, hormones, antibodies and en- the hematopoietic organs in the embryonic period. zymes (Seiverd 1972). Kidneys are also main blood production centres during the larval periods of amphibian species. Formation of Blood Thymus is the most important place of T-cell Plasma originates from the intestines and different maturation; however, the development place of B- organs of the body. Water and food substances cells is not completely clear. In adult urodeles and come from the intestines, whereas wastes, hor- anurans, the spleen is the greatest place for eryth- mones, antibodies and enzymes come from differ- rocyte production. Nevertheless, the liver also ent organs in the body (Seiverd 1972). Mature serves as a secondary organ for this activity. In blood cells are relatively short-lived; thus, they are amphibians, metamorphosis or post-hibernation continually being renewed by the differentiation hematopoiesis typically occurs in the bone mar- of stem cells produced in hematopoietic organs. The row. The bone marrow is haemopoietically active region of production blood cells depends on indi- in some salamanders (Plethodontidae) and in spe- vidual’s age. The first blood cells are produced cies Rana (Schaefer 1935, Curtis et al. 1979, Turner from mesenchymal cells in embryonic stages. In 1988). Seasonal changes could affect activity rate amphibian tadpoles, the blood cells get formed of haemopoietic organ (e.g. spleen and bone mar- Haematology of amphibians and reptiles: a review 191 row) in some species (Glomski et al. 1997). Obtaining of Blood from Reptiles Before blood is collected, the maximum blood Haemopoiesis in Reptiles volume should be determined. When reptiles are Very few studies have been made on the hae- compared with a mammalian of similar size, they mopoiesis of reptiles (Vasse & Beaupain 1981, have a lower blood volume, which constitutes 5 to Dessaurer 1970). In reptiles, the blood islets ap- 8% of the body weight. 10% of this volume can be pear to be the only erythropoietic centre during collected from healthy reptile samples (e.g. 0.5-0.8 the embryonic life. Although, like in birds, the ml in an animal that weighs 100 g). Since it is re- liver and the spleen have a hematopoietic function ported that ethylenediaminetetraacetic acid in the early periods of development, erythropoi- (EDTA) causes haemolysis particularly in turtles, esis and granulopoiesis after birth mainly occur in heparin should generally be preferred as an anti- the bone marrow. The lymphocytes that originate coagulant in reptiles (Tavares-Dias et al. 2008). from the bone marrow pass through the thymus In lizards and snakes, there are three areas for via circulation, where they get differentiated as T- blood collection by means of heparinized glass cells. It is not exactly known where the B-cells de- capillaries, namely the caudal tail vein, the postor- velop. Lymphopoiesis takes place in the bone mar- bital sinuses and the heart (MacLean et al. 1973, row, spleen and the other organs in the body. The Campbell 2006, Sykes & Klaphake 2008, Heatley & mucosa-associated lymphoid tissue (MALT) is lo- Johnson 2009). There are numerous areas for blood calized under the mucous membranes of the di- collection in turtles, including the heart and jugu- gestive, respiratory and urogenital organs. lar, subcarapacial, femoral, branchial and occipital sinuses. In crocodiles, the ventral tail vein and the supravertebral sinus are the most common areas, 2. Obtaining of Blood from which blood is collected. for Hematological Studies Obtaining the Bone Marrow Obtaining of Blood from Amphibians The bone marrow can be obtained from the femur Depending on species, season and health condi- or tibia in amphibians and most reptile species, tion, many factors affect the volume of blood cir- like in mammalians (Selleri & Hernandez Divers culating in amphibians. The blood volume was re- 2006). In turtles, the bone marrow can be obtained ported to constitute 25% of the body weight in from the area between external and internal bone caecilians but 10% of the body weight in salaman- plates (Frye 1991). In snakes, rib biopsy is the most ders and tailless frogs. realistic method to obtain the bone marrow. Blood can be collected from many areas in amphibians (Hutchison & Szarski 1965, Szarski & Czopek 1966, Allender & Fry 2008, Heatley & 3. Procedures for Counting Erythrocytes, Leuko- Johnson 2009). In most salamander and anuran cytes and Thrombocytes species, blood can be collected from the ventral abdominal vein, lingual plexus, femoral vein, fa- Blood Cell Counts cial (maxillary)/musculo-cutaneous vein, and the The erythrocytes were counted using a Neubauer ventricle of the heart (Campbell 2004, Forzán et al. haemocytometer. Standard Hayem’s or Natt and 2012). In salamanders, blood can be collected from Herrick’s (1952) solution was most used as a dilu- the ventral tail vein. Amphibians generally have tion solution for erythrocytes. The results are ex- lymphatic vessels that are widely found along the pressed as the number of erythrocytes in 1mm3 of blood vessels. The contamination of blood samples blood. with lymph might cause erroneous results. There- The total leukocyte and thrombocyte count fore, attention should be paid to lymphatic vessels acquisition for amphibian and reptilian species when collecting blood. Some researchers recom- was precluded by several factors, especially by the mend collecting blood with heparinized microhe- aggregation (Tavares-Dias et al. 2008). The leuko- matocrit tubes. The blood samples should be taken cyte count could be performed by indirect or di- to an anticoagulant tube immediately after they rect method (Campbell, 2004). The direct count us- have been collected. Information about the animal ing Natt & Herrick’s (1952) solution and with no and the date should be labeled onto the slides and dye (Hawkey 1988) or an indirect method called as blood tubes. the eosinophil pipette using 0.1% phloxine B dye 192 H. Arikan & K. Çiçek (Campbell 2004) for total leukocytes. tiles should be performed without any treatment The leukocytes were generally well counted with any anticoagulant following blood collection. by using the method of Jerrett and Mays (1973), a After blood has been dropped onto a slide, it is modified application of Blain’s method, or Natt smeared and left in order for it to dry in air and Herrick’s (1952) and with a Neubauer haemo- (Seiverd 1972). cytometer. According to Jerrett & Mays (1973), Blood smears are stained with polychrome neutral red, diluted with 0.007 of physiological stains which are the mixtures of acidic and basic water at a ratio of 1:5000, and 12% formalin mix- stains. The original polychrome stain was in- ture that was prepared again with 0.007 of physio- vented by Russian scientist Romanowsky. Many logical water were mixed at a ratio of 1:1. Natt & modifications of the Romanowsky’s stain include Herrick solution (1952) contain 3.88 g of sodium Wright’s, Giemsa, Jenner’s, May-Grünwaldand chloride, 2.50 g of sodium sulfate, 1.74 g of sodium May-Grünwald-Giemsa stains. The most popular phosphate, 0.25 g of potassium phosphate, 7.5 ml of them is the Wright’s and May-Grünwald- of formalin (37%), 0.10 g of methyl violet. The mix- Giemsa. ture dilutes to 1000 ml and filter.
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