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Chapter 5 5.1 Introduction a Fish Can Exchange Gases Effectively by The Chapter 5 5.1 Introduction A fish can exchange gases effectively by the indirect contact of blood with water in its gills. The excellent mechanism of fish gill has been studied in the fields of biology, marine zoology, and chemical engineering. Elucidation of the mechanism of a fish gill provides a hint on further improvement of artificial gill. Yamamoto et al. [1] and Schumann and Piiper [2,3] observed the change in oxygen consumption with varying the activity level of fish. Matsuda and Sakai have evaluated the oxygen transfer rate of fish gills by computer simulation analysis [4,5]. They demonstrate that the biological membrane is the rate-determining for oxygen transfer through the secondary lamellae. This is because the blood and water channels are very narrow, and these narrow channels reduce gas transfer resistances in blood and water. They also suggested that the optimal gas exchange module for membrane type artificial gill from the analysis results of computer simulation. On the other hand, the rate-determining step in artificial gill is located in the oxygen uptake device as shown in chapters 3 and 4. Thus, the oxygen uptake should be enhanced in order to improve the artificial gill. In the artificial gill, oxygen is taken up from water to the oxygen carrier solution through a gas-permeable membrane in the artificial gill, in the same way that oxygen is taken up from water to blood through a biological membrane in a biological gill. Elucidation of the oxygen uptake mechanisms of the biological gill would provides a hint on further improvement of oxygen uptake in the artificial gill. In this chapter, the oxygen transfer performance of the artificial gill were evaluated together with that of the biological gill in terms of oxygen flux, Reynolds number of water, oxygen partial pressure difference between water 131 Oxygen uptake efficiency of biological and artificial gills and blood (or oxygen carrier solution), and oxygen uptake efficiency. Advantageous for oxygen uptake in biological gill were found, and the guidelines for the further improvement of artificial gills was presented. 132 Chapter 5 5.1.1 Previous studies on fish gill Gill consists of gill arch functioning as a pillar, gill filaments standing on the gill arch like a shelf and secondary lamellae projecting numerously from the gill filaments. Inspired water from mouth is made to flow among the secondary lamellae and gill filaments. Blood channels are located in the secondary lamella, and they are separated from water by secondary lamella membrane composed of epithelium, basement membrane and pillar cell. Blood and water flow counter-currently, and gas exchange occurs. A lot of morphological information of fish gill has been obtained by many investigators up to now, and oxygen transfer mechanism of fish gill has been clarified by several researchers. Especially, the secondary lamella model established by Matsuda and Sakai, provides valuable information for designing more efficient artificial gill [4,5]. 133 Oxygen uptake efficiency of biological and artificial gills 5.1.1.1 Morphology of fish gill and water ventilation system Teleosts have five pairs of gill arches and many gill filaments standing on the front four-gill arches like a shelf. The ends of gill filaments are located in the opercular cavity. The outside of the opercular cavity is covered with an open-and-shut operculum and opened to the exterior through a gill opening (Fig .5.1). The end of gill filaments is contacted closely to the next gill filament, which produces high resistance to water flow. Hence, inspired water from mouth is made to flow through a channel between two gill filaments which has lower resistance to water flow, and effused to the opercular cavity [6-10] (Fig. 5.2). Numerous secondary lamellae are projected from both sides of the gill filaments at the right angle to them. Very narrow blood channels are located in the secondary lamella where only one blood cell can pass through (Fig. 5.3). The blood channels are separated from water by secondary lamella membrane composed of epithelium, basement membrane and pillar cell [10-16] (Fig. 5.4, Fig. 5.5). Blood is introduced to secondary lamella blood channel from afferent arch artery through afferent filamental artery. Blood and water flow counter-currently, and gas exchange occurs [10-12] (Fig. 5.6, Fig.5.7). Large partial pressure difference between water and blood caused by the counter-current produces the effective gas exchange [2,17-18]. 134 Chapter 5 Fig. 5.1 Diagram showing, in horizontal section, Fig. 5.2 Schematic diagram of piles of gill the position of the gill arches and gill filaments filaments standing on two gill arches [11,19]; and water flow: (A) Buccal cavity, (B) Buccal The end of gill filament is contacted closely to cavit y, (C) Gill arch, (D) Piles of gill filaments, the next gill filament. Water flow between slits (E) Opercul ar cavity, (F) Operculum of two gill filaments Fig. 5.3 Diagram of a portion from a gill arch of a teleost fish [20]: direction of water and blood flow are indicated by dashed and full allows, respectively 135 Oxygen uptake efficiency of biological and artificial gills Fig. 5.4 Cross-sectional view of secondary lamellae: Fig. 5.5 Electromicrograph of a secondary lamella A: cross-section of several blood channels, B: blood channel of carp (Cyprinus carpio) [19,20]: enlargement of blood channel, R: erythrocyte, P: E: epithelium, P: pillar cell, R: erythrocyte Pillar c ell, E: epithelium, N: nucleus [20] Fig. 5.6 Scanning electron micrograph of Fig. 5.7 Flow of water and blood in an a pla stic cast of a trout gill filament, showing elasmobranch fish: As in teleost fish, the flow several lamellae: Magnification 160x [20] of blood is in the opposite direction to that of the water [20]. 136 Chapter 5 5.1.1.2 Gill surface area Gill surface area per fish body weight is different for fish species. Higher values are obtained for fast and active wandering fish and lower values for elasmobranchs, freshwater fish or inactive fish [16,21]. The gill surface area is depended on fish body weight of the same kind, and the relationship between gill surface area A (mm2) and body weight W (g) is represented by the following equation [9-10]. Parameters a and m are shown in Table 5.1 for several fish species [10]. A = aW m (5.1) The gill surface area is strongly dependent on the number of gill filaments and secondary lamella density. Table 5.1 Parameters a and m in equation (5.1) representing the relationship between gill surface area and fisg body weight 137 Oxygen uptake efficiency of biological and artificial gills 5.1.1.3 Property of fish blood Fish blood consists of blood cells and plasma similar to other vertebrate animals, and the blood cells are composed of erythrocytes, leukocytes and thrombocytes. Erythrocyte is a round or ellipse shaped cell and contains dense hemoglobin. Molecular weight of fish hemoglobin is ranged from 60,000 to 70,000, which are similar to mammalian hemoglobin. There are some kinds of hemoglobin in a fish species, which are divided into several components by electrophoresis. Therefore, oxygen binding affinity and effect of carbon dioxide are different for each kinds of hemoglobin [22]. The size of erythrocyte for active fish is smaller (major axis: 7.2-14 mm, minor axis: 6.6-10 mm) than inactive fish (9.4-16 mm and 6.9-11 mm).The number of erythrocytes is 3.0-3.9 million /mm3 for active fish and 1.4-3.0 million /mm3 for inactive fish. One gram of fish hemoglobin combines with 1.48 cm3 (STP) of oxygen. The concentrations of hemoglobin for carp (Cyprinus carpio) and dogfish (Scyliorhinus stellais) are 81 mg/cm3 and 36 mg/cm3, respectively [22]. Fig. 5.8 shows the oxyhemoglobin dissociation curve of carp blood [22], and Table 5.3 shows the half saturation p50 (mmHg) (the oxygen partial pressure when degree of oxygen saturation of hemoglobin is 0.5). The half saturation p50 for fish blood is ranged from 5 to 20 mmHg [22], which is lower than that of human beings (26 mmHg) [23]. 138 Chapter 5 Fig. 5.8 Oxyhemoglobin dissociation curves of carp blood [22] Table 5.2 Half saturation of hemoglobin for fish blood [4] p50 Oxygen cap. Temperature Fish species (mmHg) (vol%) (K) Ameiurus nebulosus 1.4 13.3 288 Cyprinus carpio 5 12.5 288 Scyliorhinus stellais 15.8 5.3 290 Amia calva 4.0 11.8 288 Salmo gairdnerii 18 13.8 288 Salvelinus fontinalis 13 11.7 288 Scomber scombrus 16 15.8 293 Human being (male) 26 19.8 310 139 Oxygen uptake efficiency of biological and artificial gills 5.1.4 Oxygen consumption of fish Several researchers revealed that oxygen consumption rate of fish varies with the activity level of them [1,24,25]. Yamamoto et al. measured the oxygen consumption rate of a Carp with various ventilation rates by the respire-chamber method as shown Fig.5.9. The results of this study provided a lot of valuable information. Actually, Matsuda and Sakai used the experimental results to establish the secondary lamella model [4,5]. In this chapter, the oxygen consumption rate that was measured Yamamoto et al., was also used to evaluation of oxygen uptake performance of the present artificial gill. Fig. 5.9 Schematic of apparatus used Yamamoto et al. for measuring oxygen consumption rate of carp [1] 140 Chapter 5 5.2 Experimental and Theoretical section 5.2.1 Oxygen uptake of biological and artificial gills Fish take up oxygen from water into their blood through a secondary lamella of the gill.
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