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Research Article Histological Differences in Axial Musculature

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Iran. J. Ichthyol. (June 2020), 7(2): 136-147 Received: February 25, 2019 © 2020 Iranian Society of Ichthyology Accepted: June 27, 2020 P-ISSN: 2383-1561; E-ISSN: 2383-0964 http://www.ijichthyol.org Research Article Histological differences in axial musculature between relative species of the genera Phoxinus and Rhynchocypris (Actinopterygii: Cypriniformes: Leuciscidae) Nikita O. YABLOKOV1,2, Ivan V. ZUEV*2 1Scientific Research Institute of Ecology of Fishery Reservoirs, Krasnoyarsk, Russia. 2Siberian Federal University, Krasnoyarsk, Russia. *Email: [email protected] Abstract: Comparative studies of axial musculature in Phoxinus phoxinus and three species of the genus Rhynchocypris were conducted to analyze locomotor abilities and biotopic preferences of widespread Asian minnows. The ratio between the areas occupied by the red and white muscles, as well as the fiber diameter, was measured on histological sections. Significant between-species differences were found in the size of the both type of fibers. Mean diameter of white fibers varied from 50.7 to 69.3μm, the extreme values (min-max) of this parameter were found in typical lotic species, R. lagowskii and P. phoxinus, respectively. Mean diameter of red fibers varied from 26.4 to 33.5μm, and also does not have a clear link to the type of habitat. On the contrary, the distribution of different types of fibres was in good agreement with the data on preferred biotopes of studied species, divided into two groups. Mean proportion of red fibres in caudal peduncle in lotic P. phoxinus and R. lagowskii (9.1- 10.2%) was more than twice as high as that in lentic R. percnurus and R. czekanowskii (3.8- 4.3%). The low proportion of red muscles in R. percnurus and R. czekanowskii, in addition to poor mobility, may also indicate adaptation to wintering under hypoxic conditions. In general, the ratio of red to white muscles provides additional information for predicting the distribution of minnows in freshwaters. Keywords: Minnows, Fibre size, Red muscles, White muscles, Preferred biotopes, Northern Asia. Citation: Yablokov, N.O. & Zuev, I.V. 2020. Histological differences in axial musculature between relative species of the genera Phoxinus and Rhynchocypris (Actinopterygii: Cypriniformes: Leuciscidae). Iranian Journal of Ichthyology 7(2): 136-147. Introduction Such uneven distribution of different minnow species Minnows of the genera Phoxinus and Rhynchocypris from the center of the origin cannot be fully are widespread in the Palearctic, which is inhabited explained by either theory of ocean level fluctuation by 7-9 species of Phoxinus and 5-6 species of in the quaternary period (Lindberg 1972) or Rhynchocypris (Sakai et al. 2006; Kottelat & Freyhof molecular phylogenetic data (e.g. Imoto et al. 2013; 2007; Schönhuth et al. 2018). Ranges of the Eurasian Schönhuth et al. 2018). An alternative approach to minnow, P. phoxinus and most of the Rhynchocypris the analysis of the modern ranges of species, as well species overlap in the Russian Far East which is as the forecast of their modification due climate considered to be the center of speciation of change, is to study their certain ecological niches (Yu Leuciscidae (Ito et al. 2002; Sakai et al. 2006; Perea et al. 2013). et al. 2010). The area outside the region to the west is Fish morphology is a basic and simple tool for inhabited only by R. czekanowskii and R. lagowskii determining the ecological niches in closely related (to the East Siberia), R. percnurus (to the East species (Gatz Jr 1981; Douglas & Matthews 1992). Europe) and the Eurasian minnow (to the Iberian However, in the case of minnows from the genus Peninsula) (Berg 1948; Kottelat & Freyhof 2007). Rhynchocypris, which are very similar in 136 Yablokov and Zuev - Histological differences in axial musculature appearance, the external morphology for the given purpose is not effective. At the same time, external similarity does not exclude possible differences in the structure of the axial muscles. Axial musculature of many fishes is distinctively divided at least into two types of muscle: red oxidative and white glycolytic. Red muscles are powered by oxidative phosphorylation, whereas white muscles are largely powered by anaerobic utilization of phosphocreatine, ATP and glycogen (Svendsen et al. 2015). Although any type of swimming is a complex combination of using of different muscles, red fibres are more active during sustained swimming, whereas white fibres are used at high swimming speed (Sanger & Stoiber 2001). Thereby, the ratio between these types of muscles correlates with capable of active and long- term movements and partly can predict the ecological requirements of fishes (Boddeke et al. 1959; Slijper 1963; Langerhans 2008). By now the anatomical Fig.1. Map of sampling sites in the Northern Asia. features of different types of muscles and their proportions are not known for all taxonomic groups 2016 in different water bodies of the Yenisei, Pyasina of fishes. Relatively many data have been published and Amur River basins (Fig. 1, Table 1). All the on marine species (Greeк-Walker & Pull 1975; specimens were killed by overdose of clove oil Mosse & Hudson 1977), while freshwater fishes are following the guidelines given by the Institutional poorly studied in this respect. Ethical Committee and were preserved in 4% neutral In this paper, we compare the ratio of red and formalin. Before sectioning, total length (TL, mm) white muscles and the size of their fibers among four was determined. related species of genera Phoxinus and Muscle sectioning: Fishes were cut into four sections Rhynchocypris from Siberia and the Russian Far with a blade (Zhang et al. 1996; Drazen et al. 2013). East. The aim of the comparison was to test a link Slice A was at the back of the base of the pectoral fin, between studied parameters and the type of preferred slice B was at the beginning of the base of the dorsal biotope and other environmental factors. fin, and slice C was at the end of the base of the anal fin (Fig. 2). Each section, except for the head, was Materials and Methods mounted under stereomicroscope Micromed MC2 Sample collection: The ratio between different types Zoom 2CR; the anterior part of each section was of axial muscles and their fibre diameter were photographed by digital camera ToupCam 5.1 estimated for the four species of the genera Phoxinus (ToupTek, China). For slices exceeding the field of and Rhynchocypris: Eurasian minnow P. phoxinus view of the microscope, a series of four photographs (Linnaeus, 1758), lake minnow R. percnurus (Pallas, which included layers of epaxial and hypaxial 1814), Czekanowskii minnow R. czekanowskii muscles of the right and left parts of the slice was (Dybowski, 1869) and Amur minnow R. lagowskii taken. Red musculature was considered as triangular (Dybowski, 1869). Fishes were caught by patches of dark color, located laterally between the electrofishing during the summer months of 2013- layers of epaxial and hypaxial white muscle tissue 137 Iran. J. Ichthyol. (June 2020), 7(2): 136-147 Table 1. Sample sites and samples description. River basin Geographical Type of Sampling № Species TL ± SE, mm N coordinates water body period 57°27'08.5" N, August 73.6 ± 4.2 1 Yenisei Lake 5 Rhynchocypris 93°11'03.6" E 2016 62.0 –85.0 czekanowskii 69°22'48.1"N July 90.0 ± 5.4 2 Pyasina Lake 5 88°11'41.2"E 2013 71.0 – 108.0 Rhynchocypris 56°34'57.3"N, August 60.9 ± 0.7 3 Yenisei Lake 10 percnurus 93°43'23.6"E 2016 56.0 – 64.0 Rhynchocypris 57°27'08.5"N, June 106.5± 0.4 4 Amur Reservoir 10 lagowskii 93°11'03.6"E 2014 84.0 –122.0 Phoxinus 57°27'08.5"N, July 75.9 ± 2.1 5 Yenisei River 10 phoxinus 93°11'03.6"E 2015 63.0 –87.0 Table 2. Duration of exposure of minnow’s muscle samples at the Microm STP 120 automated station. № Reagent Exposure, min 1 Ethanol 70° 30 2 Ethanol 80° 30 3 Ethanol 90° 30 4 Ethanol 96° 35 5 Ethanol 96° 35 6 Ethanol 100° 35 7 Ethanol 100° 35 8 Xylol 45 9 Xylol 45 10 Paraffin 60 11 Paraffin 80 Microm STP 120 (Table 2). Paraffin was poured into samples using a filling system Thermo Scientific Fig.2. Position of slices and distribution of white (1) Microm EC 350-1; sections were prepared using a and red (2) muscles. semi-automatic microtome Microm HM 440E. (Fig. 2). The area occupied by the red and white Thickness of histological sections was 10μm. muscles was measured in photographs in the ImageJ Obtained histological sections were viewed under 1.51r graphics editor (NIH, USA). Proportion of red a Leica DMLC light microscope equipped with a muscles was estimated as a percentage of the total digital camera (Leica DC100). Photographing of area occupied by the muscles in the cross section for sections was performed under different each slice. magnification. The shortest fiber diameter, calculated Histological preparation. About 1cm3 size blocks perpendicular to the longest diameter at its mid-point, were extracted from section B-C (Fig. 2). For the was measured by digital image using ImageJ 1.51r histology examination, samples were dehydrated, software (NIH, USA). For each examined fish, the cleared with xylol, then embedded in paraffin and diameters of 50 red and 50 white fibres were stained with hematoxylin and Ehrlich’s eosin measured at random. (Mumford 2004). Dehydration was carried out by Statistical analyses. Statistical analyses were incubation in ascending concentrations of alcohols performed with PAST 3.17 (Hammer et al. 2001). (xylol and ethanol) using an automatic station Fibre diameter and red muscle proportion is 138 Yablokov and Zuev - Histological differences in axial musculature Table 3.
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