Ahe 1014

Anatomia Histologia Embryologia
1 ORIGINAL ARTICLE
2
3 Study of Epidermis Development in Sturgeon (Acipenser
4
5
persicus) Larvae
6 Z. Saadatfar1, D. Shahsavani2 and F. S. Fatemi3
7
Addresses of authors: 1 Department of Anatomical Science, School of Veterinary Medicine, Ferdowsi University of Mashhad, Mashhad, Iran;
8 2
Department of Food Hygiene and Aquaculture, School of Veterinary Medicine, Ferdowsi University of Mashhad, Mashhad, Iran;
9 3
Department of Environmental Technology, Motahar Institute
10
11
12
13
14 *Correspondence: Summary
15 Tel.: 91775-1793;
16 e-mail: Saadatfar.z@gmail.com; Fish skin is essential for survival, maintenance of body shape, and protection 1
17 saadat_histo@yahoo.com against the shock and infection. During week 1 of sturgeon larval development,
18 the epidermis is thin and not differentiated in various layers yet, but by week
With 10 figures 4–6 the thickness increases and various layers appear, depending upon the
19
20 region of the body. Mucous cells differentiate early in development on the sur-
Received August 2009; accepted for
21 publication May 2010
face of epithelium and contain acid and neutral mucopolysaccharides. Primor-
22 dial sensory buds are visible within the epithelia of the skin of the head in
23 doi: 10.1111/j.1439-0264.2010.01014.x week 1 larvae, and become numerous during later larval development. Club
24 cells are specialized epidermal cells that have an immune function and appear
25 in the middle layer of the head and trunk epidermis on week 4.
26
27
28 The skin has several important functions: it provides
Introduction
29 physical protection for the body, improves hydrodynam-
30 Fish skin is a vast and important organ which has a ics, helps in maintaining the osmotic balance, contains
31 major role in adaptation of fish to the environment and sensory receptors, and in some fishes also takes part in
32 in the protection against infections. It differs from the respiration. The skin also plays a substantial role in swim-
33 exposed skin of other vertebrates by the presence of living ming mechanics as it contributes to the overall skeletal
34 superficial epidermal cells which are in direct contact with support, and muscle force must be transmitted down the
35 the aqueous environment, and by a superficial mucous body through the skin. Furthermore, fish skin contains
36 layer instead of a keratin layer (Dominique et al., 2004). mucous glands which aid in maintaining the water bal-
37 The Persian sturgeon, Acipenser persicus, is widely distrib- ance, and enzymes and antibodies, which inhibit entry of
38 uted in the southern basin of the Caspian Sea and is a bacteria that are extremely numerous on the skin surface
39 native fish species of the Iranian coasts. It is one of the (Rottmann et al., 1992). In a number of fish species, the
40 most valuable species of Acipenseridae, but is in danger skin also offers protection through control of coloration
41 of extinction (Bauer et al., 2002). During the past century by expansion and contraction of the chromatophores,
42 sturgeon population has diminished by overfishing, lack and in some species epidermal respiration occurs by the
43 of management control and by industrial and domestic exchange of oxygen and carbon dioxide through the
44 pollution. This species is used for commercial food prod- numerous blood vessels near the skin surface.
45 ucts such as caviar and smoked sturgeon meat or soup, The structure, organization and function of fish epider-
46 while sturgeon skin provides valuable leather material for mis have interested researchers for a long time. Fish skin
47 clothing, shoes and handbags. Sturgeon production is is composed of three compartments: epidermis, dermis
48 considered as a business with great economic potential, and hypodermis. The epidermis comprises only living
49 but fingerling production in hatcheries is often accompa- cells, in contrast to that of terrestrial vertebrates which
50 nied by the risk of infection. Therefore, in-depth study of are covered by an outer layer of dead and keratinized
51 the integument, which forms the outer protective barrier cells. Such a layer is uncommon in fish, and occurs only
52 against aquatic environment, can increase our knowledge in specific sites that are subjected to abrasion such as
53 to manage and reduce the risk of diseases in aquaculture. adhesive organs, lips and pads, and on the epidermal
ª 2010 Blackwell Verlag GmbH • Anat. Histol. Embryol. 1
A H E
Journal Name
1 0 1 4
Manuscript No. B Dispatch: 14.6.10
Author Received:
Journal: AHE CE: Shaahul
No. of pages: 6 PE: Gayathri
xxxxx Z. Saadatfar, D. Shahsavani and F.S. Fatemi 21
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1 surface of some species capable of emerging from the
2 water (Whitear, 1977).
3 There is little information regarding the development
4 of fish skin, particularly in Acipenseridae. Their skin has
5 special morphological, histological and physiological char-
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6 acteristics, such as a beautiful appearance, a specific
7 arrangement of the scales and a well-balanced thickness
8 and flexibility that makes it suitable for production of
9 high quality leather and related products. The present his-
10 tological and histochemical study of the structure of fish
11 skin and its mucous components during subsequent larval
12 stages will help to understand the process of skin devel-
13 opment, and also to diagnose and treat fish skin diseases.
14
15
Material and Methods
16
17 Acipenser larvae of 1–44 days post-hatching (dph) were
18 obtained from Shahid Marjani Propagation Center in Fig. 1. Histological section of the trunk skin of 2 days old larvae 11
19 Gorgon. The specimens were kept in fibreglass tanks to (PAS). mu: mucous cell me: mesenchyme.
20 21 dph and were transferred to pond until the age of
21 44 days (fingerling stage). About five fish of each larval

22 stage were studied. The time of sampling was every sec-
23 ond day. Specimens were fixed in 10% neutral buffered
24 formaldehyde and transferred to the laboratory. These lar-
25 vae were later dehydrated through a standard ethanol ser-
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26 ies to 100%, cleared in xylene, embedded in wax, cut into
27 7-lm-thick sections using a microtome, deparaffinized
28 and stained with haematoxylin & eosin for general histol-
29 ogy. Mucous substances were demonstrated by alcian blue
30 solution (AB) at pH 1 and pH 2.5 (Steedman, 1950; Lev
31 and Spicer, 1964), periodic acid - Schiff (PAS) method
32 and AB-PAS staining (Lillie and Greco, 1974), whereafter
33 they were studied using light microscopy.
34
35
Results
36
37 Week 1
38 The epidermis was evident from 1 dph. The number of
39 epidermal layers altered from 2 (1 dph) to 10 (7 dph) in
40 function of time. Epidermal thickness varied at different
41 body regions (Fig. 1), being thicker at the head region
42 compared with the trunk (Fig. 2). Large and light round
43 cells were present from the onset of larval development
Fig. 2. Histological section of head skin in 7 day old larvae (PAS). mu: 12
44 and were considered as mucus producing cells. Some of
mucous cell sb: sensory bud me: mesenchyme.
45 the round cells had a positive reaction to PAS (Fig. 2)
46 and AB (pH 1 and 2.5). At the end of week 1, sensory
47 buds were formed on the epidermis of the head and enhanced. The sensory buds had also increased in size
48 assumed circular forms. and number. Mucous cells constituted the majority of
49 cells in the epidermis and were scattered over the trunk,
50 occupying the whole epidermal thickness (Fig. 3). Some
Week 2
51 of these cells had a positive reaction to PAS and AB (pH
52 The thickness of the epidermis and the size of both the 1 and 2.5), while other cells did not show any response
53 epidermal cells and the light round cells were clearly (Fig. 4, 5, 6).
2 ª 2010 Blackwell Verlag GmbH • Anat. Histol. Embryol.

Z. Saadatfar, D. Shahsavani and F.S. Fatemi xxxxx 21

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2
3
4
5
6
7
8
9
10
11
12
13
14
15
16 Fig. 3. Histological section of the head skin of 2 weeks old larvae 13
17 (AB). mu: mucous cell sb: sensory bud ch: chromatophore. Fig. 5. Histological section of the trunk skin of 2 weeks old larvae 15
18 (PAS). mu: mucous cell co: collagen.
19
20
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21
22
23
24
25
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31
32
33
34
35
36
Fig. 6. Histological section of the trunk skin of 2 weeks old larvae 16
37 (AB-PAS). mu: mucous cell co: collagen ch: chromatophore.
38
39 Fig. 4. Histological section of the trunk skin of 2 weeks old larvae 14
40 (AB). mu: mucous cell co: collagen. prised up to 25 layers. At the end of the fifth week, an
41 epidermis consisting of three layers was evident in some
42 parts of the head. The superficial layer consisted of cuboi-
Week 3
43 dal to squamous epithelial cells and round mucus cells.
44 No specific changes were observed during this develop- The middle layer comprised large globular cells, whereas
45 mental stage. However, an increase in epidermal thickness the basal layer was composed of columnar cells (Fig. 8).
46 and size of cells and sensory buds was detected. Some Two types of circular cells were found in the epithelium.
47 mucus cells showed a more intense reaction to histo- The smaller cell type was located more superficially, and
48 chemical staining (Fig. 7). showed a positive reaction to PAS and AB staining which
49 became more intense as the larvae grew (Fig. 9). The lar-
50 ger circular cells were situated in the middle layer of the
Weeks 4, 5 and 6
51 epidermis. They presented no positive response to any
52 At this phase the epidermal thickness increased manifestly histochemical staining, and were assumed to be club or
53 and the epithelial cells of certain regions of the head com- alarm cells (Fig. 9, 10). The number and size of sensory


1
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2
3
4
5
6
7
8
9
10
11
12
13
14
15
17 (H&E). mu: mucous cell sb: sensory bud me: mesenchyme.
18
19
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20
21
22
23
24 Fig. 9. Histological section of the head skin of 6 weeks old larvae (AB 19
-PAS). mu: mucous cell cl:club cell sb:sensory bud. ch: choromato-
25
phore me: mesenchyme.
26
27
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28

29
30
31
32
33
34
36 (H&E). bl: basal layer ml: middle layer sl: superficial layer. mu: mucous
cell cl:club cell sq:squamous cell.
37
38
39 buds in the head region increased during larval develop-
40 ment. They consisted of up to 50 cells gathered around
41 the glandular opening, but had a negative reaction to
42 PAS and AB staining at all stages.
43 Fig. 10. Histological section of the trunk skin of 6 weeks old larvae 20
44 (H&E). mu: mucous cell cl:club cell sq:squamous cell me: mesen-
Discussion chyme.
45
46 Several studies have been performed on adult teleost skin
47 (Kapoor, 1966; Hawkes, 1974; Weisel, 1978; Hebrank and tion of the mucopolysaccharides in the stages of skin
48 Hebrank, 1986; Fishelson, 1988; Mittal and Datta Munshi, development in Acipenser persicus.
49 2 2005), but little information is available about the embry- The superficial layer of the epidermis of this species
50 onic and larval stages (Le Guellec et al., 2004; Foscarini, was composed of living cells. Keratinization of the skin is
51 2006; Webb and Kimelman, 2008). As information on uncommon in fish, except for a number of species such
52 sturgeon skin is lacking, the aim of this study was to elu- as Solea senegalensis in which the head region is described
53 cidate the differentiation of the epidermis and the forma- as a specific site of keratinization (Whitear, 1977; Arellano

Z. Saadatfar, D. Shahsavani and F.S. Fatemi xxxxx 21
1 et al., 2004). In most fish species, however, the skin is Differentiating club or alarm cells were found when the 6
2 devoid of a protective layer of keratin, while the functions larvae are transferred from tank to pond (week 4). These
3 of the latter are in part compensated by the secretion of cells were observed in the middle layer of the head epi-
4 3 mucus (Carlos, 1994). dermis and at the surface of trunk epidermis and had a
5 Apparent changes in epithelial thickness were observed, negative response to histochemical staining. An earlier
6 varying from 2 to 25 layers in the period investigated. study suggested that these cells have an immunological
7 According to the previous findings, epidermal thickness function protecting against general injury, parasites and
8 differs according to fish species, age and body sites pathogens, while as a secondary role they may have an
9 (Campinho, 2007; Kim et al., 2008). Thickness can serve alarming function (Chivers and Wisenden, 2007).
10 as a protection against physical injury and to establish a The sensory buds in Acipenser persicus were observed in
11 first line defence to mechanical stress. The epithelium the head, lip, branchial and facial skin. The shape and size
12 may form a selective permeability barrier for water and of the sensory buds varied in different parts of the head
13 ions, while environmental factors may determine and region and contained species-specific epithelial cells with
14 affect epidermal thickness (Wendelaar and Meis, 1981). various cellular components which need further investiga-
15 Additionally, the skin plays a major mechanical role tion by electron microscopy. Because these structures are
16 in swimming and behavioural patterns (Hebrank and confined to the epidermis of the head region, they could
17 4 Hebrank, 1986). be sensory organs of the olfactory system or taste buds. It
18 The epithelial cells of the head epidermis were more is postulated that the differences in these organs are
19 differentiated at the end of the developmental period. related to ecomorphological adaptation (Fishelson and
20 They formed a stratified epithelium consisting of three Delavea, 2004; Ohkubo et al., 2005; Cinar et al., 2007). 7
21 layers, viz. a superficial, middle and basal layer. This In conclusion, many of the present findings in stur-
22 observation is similar to the findings in other fish species geons are in agreement with the previous reports on epi-
23 such as torrent catfish (Liobagrus mediadiposalis) (Park dermis of fish skin. From the first day after hatching,
24 5 et al., 2003a), Senegal sole (Arellano et al., 2004), Korean different mechanisms are triggered to develop a func-
25 loach (Iksookimia koreensis) (Park, 2002) and eel goby tional stratified epidermis for the protection of the larvae
26 (Odontamblyopus lacepedii) (Park et al., 2003b). This against various external factors. Further electron micro-
27 study suggests that differentiation of cells proceeds with scopic study might lead to a precise knowledge of epider-
28 transfer of larvae from tanks to ponds. mal cell morphology and function during the process of
29 The upper cell layer consisted of cuboidal and squa- skin development.
30 mous cells, of which some showed signs of cellular degen-
31 eration. The cytoplasm was darkly stained and nuclei
Acknowledgements
32 were indistinguishable. Some of these cells were detached
33 from the surface and could most probably be replaced by The authors are sincerely grateful to Dr.Khayatzadeh for
34 the deep cells. Mucous cells scattered in the superficial her guidance through the study.
35 layer can ensure protection against bacteria, fungi and
36 parasites.
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38 cells, including a variable number of mucous cells, club Arellano, J., M. V. Storch, and C. Sarasquet, 2004: Ultra-
39 cells and or undifferentiated cells. The histochemical tech- structural and histochemical study on gills and skin of
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48 AB and also to combined AB-PAS staining, whereas other
scale variation in epidermal thickness, distribution, and size
49 cells did not have any response to histochemical staining.
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Anatomia Histologia Embryologia

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