CC BY
2
ФИЗИОЛОГИЯ И БИОХИМИЯ ГИДРОБИОНТОВ
PHYSIOLOGY AND BIOCHEMISTRY OF HYDROCOLE
Original article UDK 597
https://doi.org/10.24143/2073-5529-2023-3-82-89 EDN JNTTIT
Morphophysiological and biochemical studies of Caspian atherina (Atherina boyeri caspia Risso, 1810)
Pirmurad S. Taibov1, Nyria A. Kanieva2^, Vitalii V. Barabanov3, Natalia V. Kozlova4
West-Caspian Department of the Volga-Caspian Branch Russian Federal "Research Institute of Fisheries and Oceanography ", Makhachkala, Russia
2Astrakhan State Technical University, Astrakhan, Russia, kanievana52@mailruM
3 4Volga-Caspian Branch Russian Federal "Research Institute of Fisheries and Oceanography ",
Astrakhan, Russia
Abstract. The article highlights physiological and biochemical parameters of atherina (Atherina boyeri caspia Risso, 1810) in the Middle Caspian Sea at various stages of the annual cycle. The state of the fish organism was studied in the period from February to October 2022, which characterized the pre-spawning, spawning and feeding periods. The size-weight and age parameters of atherina were determined. The concentration of the main reserve substances of the fish body - total lipids and water-soluble protein - was analyzed in the muscles of the atherina. In the pre-spawning period and at the beginning of spawning of females, the level of morphological changes in the gonads was studied. There were no found significant differences in the content of total lipids and water-soluble protein in the muscles of males and females. Patterns of changes in the amount of muscle water-soluble proteins and total lipids were recorded in the entire sample of the Caspian atherina during the pre-spawning period and at the beginning of spawning. This is typical for portioned spawning fish, in which the energy expenditure of the body is directed to the formation, maturation of gonads, the removal of sexual products and the maturation of the next portions during the spawning period. In September-October, that is, already at the stage of feeding, a multidirectional character in lipid and protein metabolism was noted in the body of atherina. According to the results of histological examination, the beginning of spawning in atherina was noted in April. Mature oocytes with resorption at an average level of 30-31% were recorded in the ovaries of portioned spawning atherina in February-April, which characterized the adaptive mechanisms of the organism in the conditions of this habitat. In September, there was a significantly positive (p < 0.05) statistical correlation between the concentration of total muscle lipids and the mass of fish (r = +0.47), which characterized a positive energy balance in the body, so necessary for the next stages of life.
Keywords: atherina, muscles, gonads, total lipids, water-soluble protein
For citation: Taibov P. S., Kanieva N. A., Barabanov V. V., Kozlova N. V. Morphophysiological and biochemical studies of Caspian atherina (Atherina boyeri caspia Risso, 1810). Vestnik of Astrakhan State Technical University. Series: Fishing Industry. 2023;3:82-89. (In Russ.). https://doi.org/10.24143/2073-5529-2023-3-82-89. EDN JNTTIT.
© Taibov P. S., Kanieva N. A., Barabanov V. V., Kozlova N. V., 2023
Научная статья
er
o с
o
о. b
Морфофизиологические и биохимические исследования каспийской атерины (Atherina boyeri caspia Risso, 1810)
Пирмурад Султанмурадович Таибов1, Нурия Абдрахимовна Каниева2ш, Виталий Викторович Барабанов3, Наталья Викторовна Козлова4
1Западно-Каспийский отдел Волжско-Каспийского филиала Всероссийского научно-исследовательского института рыбного хозяйства и океанографии, Махачкала, Россия
2Астраханский государственный технический университет, Астрахань, Россия, kanievana52@mail.ruM
3 4Волжско-Каспийский филиал Всероссийского научно-исследовательского института рыбного хозяйства и океанографии, Астрахань, Россия
Аннотация. Исследованы физиолого-биохимические показатели атерины (Atherina boyeri caspia Risso, 1810) в Среднем Каспии на различных этапах годового цикла. Состояние организма рыб изучали в период с февраля по октябрь 2022 г., что характеризовало преднерестовый, нерестовый и нагульный периоды. Определяли размерно-весовые и возрастные показатели атерины. В мышцах атерины анализировали концентрацию основных резервных веществ организма рыб - общих липидов и водорастворимого белка. В преднерестовый период и в начале нереста у самок исследовали уровень морфологических изменений гонад. В мышцах самцов и самок достоверных различий содержания общих липидов и водорастворимого белка не зафиксировано. У всей выборки каспийской атерины в преднерестовый период и на начало нереста зарегистрированы закономерности изменения количества мышечных водорастворимых белков и общих липидов. Это характерно для порционно нерестующих рыб, у которых затраты энергии организма направлены на формирование, созревание гонад, вы-мет половых продуктов и дозревание очередных порций в период нереста. В сентябре-октябре, т. е. уже на этапе нагула, в организме атерины был отмечен разнонаправленный характер в липидном и белковом обмене. По s результатам гистологического исследования начало нереста у атерины отмечено в апреле. В яичниках порционно i нерестующей атерины в феврале-апреле регистрировали зрелые ооциты с резорбцией на среднем уровне s 30-31 %, что характеризовало адаптационные механизмы организма в условиях данной среды обитания. f В сентябре отмечена достоверно положительная (p < 0,05) статистическая зависимость между концентрацией s общих мышечных липидов и массой рыб (r = +0,47), что характеризовало положительный энергетический ба- Ц-ланс в организме, так необходимый для следующих этапов жизнедеятельности. t
e
Ключевые слова: атерина, мышцы, гонады, общие липиды, водорастворимый белок n
a
Для цитирования: Таибов П. С., Каниева Н. А., Барабанов В. В., Козлова Н. В. Морфофизиологические t
и биохимические исследования каспийской атерины (Atherina boyeri caspia Risso, 1810) // Вестник Астрахан- Г
ского государственного технического университета. Серия: Рыбное хозяйство. 2023. № 3. С. 82-89. i
https://doi.org/10.24143/2073-5529-2023-3-82-89. EDN JNTTIT. О
y
i
Introduction tion and variability of its morphometric features have §
The Caspian form of atherina (Atherina boyeri cas- been studied [4-6]. a
pia Risso, 1810) is widespread throughout the Caspian However, in the literature there are only fragmen- i
Sea. It is a marine euryhaline fish species [1] that lives tary data on morphological and biochemical parame- §
in desalinated pre-estuarine areas, in river mouths [2], as ters of Caspian atherina and there is no data on its 1
well as in its most saline areas. Recently, atherina in the physiological capabilities, especially on the metabolic 3
Caspian Sea, in terms of its mass, occupies one of the processes of the body that change during the period
second place after the Caspian sprat. of individual development.
Atherina is the main object of by-catch in fishing The purpose of the work is to undertake physiologi-
Caspian sprat with fixed nets. In recent years, an in- cal and biochemical studies of atherina at various stages
crease in by-catch of atherina has been recorded in of the annual cycle in the Caspian Sea. catches of Caspian sprat [3].
There is information in the literature about the size, Research materials and methods weight and age characteristics of the spawning part The work was carried out in the West Caspian De-
of the atherina population in the modern ecological con- partment of the Volga-Caspian Branch of the Federal
ditions of the Caspian Sea. The issues of atherina nutri- State Budgetary Institution "VNIRO" (CaspNIRKH)
in the period of 2014-2021. The research was carried
о ^
s? а
X
« §
о «
s с
out in the waters of the Kizlyar Bay and the adjacent area between the Bryansk and Suyutkino spit, on the Krainovsky coast from the Suyutkino spit in the south to the northern tip of Chechen Island.
Sampling was carried out in accordance with the instructions [7] in February, March, April, July, September, and October. The mass, length and gender were determined according to the generally accepted method [8]. The total lipid content in the muscles was determined by a modified method with phosphorus-vanillin reagent according to Zollner-Kirsch [9], the Warburg and Christian spectrophotometric method was used to determine the concentration of water-soluble protein [10, 11]. The determination of the stages of maturity of the gonads (NWG) of females was
carried out by standard histological methods [12]. The results of the obtained data are presented in the form of average values and their errors (M ± m). Statistical processing was carried out, according to the modern data processing rules [13, 14].
Research results and their discussion
The average linear-weight parameters of atherina (8.4 cm and 5.5 g) in 2021 were close to the average long-term indicators for the last 5 years (2016-2020). The average Fulton's stomach fullness index was 0.944, which indicates a high feeding intensity of fish. During the study period, the predominance of females in the gender composition of the atherina population was noted - from 56 to 59% (Table 1).
Table 1
Biostatistical indexes of atherina in 2016-2021
Years Total catch, t Сatch on fishing effort, t Average length, cm Average weight, g Proportion of females, % Fulton's stomach fullness index
2016 48.4 6.0 8.2 5.2 58 0.943
2017 34.6 3.5 8.3 5.4 57 0.944
2018 60.6 5.9 8.1 5.2 59 0.978
2019 57.6 5.3 8.4 5.5 55 0.928
2020 66.1 3.7 8.3 5.4 56 0.944
2021 70.7 2.7 8.4 5.5 58 0.927
Average of2016-2021 56.3 4.5 8.3 5.4 57.2 0.944
ч о s
M
s -е-
о
-е-&
о
и к
и и
w <г
к
ы и с
ш о ю s rt H
In the course of the research the size and weight characteristics of atherina caught in different seasons
of 2022 in the Middle Caspian region were determined (Table 2).
Table 2
Mass and length of the studied individuals of atherina in the Caspian Sea in 2022
Indexes Months
February March April July September October
Mass, g 4.2 ± 0.2 4.2 ± 0.6 4.6 ± 0.4 6.5 ± 0.3 6.2 ± 0.2 6.5 ± 0.3
Length, cm 8.2 ± 0.1 9.3 ± 0.4 9.8 ± 0.3 9.8 ± 0.2 10.1 ± 0.1 10.1 ± 0.2
In a living organism there take part constant metabolic processes characterized by the synthesis and decomposition of various biochemical compounds. To obtain information about synthesis and decomposition it is necessary to study the content of biochemical substrates of the organism aimed at the vital activity of cells and tissues in different periods of the annual cycle [15, 16].
Proteins and lipids are the main material substrate in animal tissues, the change of which characterizes plastic and energy metabolism [16].
Caspian atherina caught in February-March 2022 was characterized by a pre-spawning condition in April fish spawned.
In the pre-spawning period of February-March the results of studies of the amount of total lipids and water-soluble protein in the muscles of male and female fish did not reveal statistically significant differences. At the same time, the lipid content in females averaged 3.05%, in males - 3.08%, the concentration of water-soluble protein was 55.49 and 54.76 mg/g respectively (Table 3).
Table 3
Content of biochemical parameters in the muscles of atherina in the pre-spawning period
Gender Total lipids, % Water-soluble protein, mg/g
Females 3.05 ± 0.24 55.49 ± 4.42
Males 3.08 ± 0.20 54.76 ± 2.16
In the absence of significant differences in the content of lipids and water-soluble protein in the muscles of females and males the analysis of physiological and biochemical parameters in individuals without sexual division is further presented in the work.
The lipid content in fish muscles is highly variable during the annual cycle [17-19]. The results of the study of total lipids and water-soluble protein in atherina muscle tissue from February to October are
shown in Fig. 1.
a*
o
<
5.30
4.30
3.30
2.30
70.00
60.00
50.00
40.00 S
30.00
20.00 Ä
10.00
00.00
>
February March April - Total lipids, %
July September October
- Water-soluble protein, mg/g
Fig. 1. Dynamics of the amount of muscle protein and total lipids in the body of atherina from February to October 2022
In April there were noted fish that had already swept out the first portion of sexual products. The content of total lipids in the muscles of atherina was registered in the amount of 3.11%. Thus, in the period from March to April an increase in total lipids in the muscles was noted, since the main forming processes in the gonads had already ended, and during the transition of the gon-ads from stage IV to stage V of gonad maturation (SGM), mainly only their watering occurs [17]. During the spawning period the mobilization of lipids of atheri-na occurs, i. e. their accumulation. According to the literature data [4], the sexually mature atherina living in the Middle Caspian Sea does not have empty intestines in the spring and summer periods, i. e., a stable energy balance is established. Obviously, this is due to the process of shedding of roe and milt, which require significant energy consumption, as well as an increase in the mobility of fish [17]. In the Middle Caspian Sea atherina spawning begins in spring and lasts almost all summer [4]. In July the total lipid content in the muscles of atherina was at the level of April parameters - 3.11%.
In September-October the accumulation of lipid reserves occurs in the muscle tissue of atherina after spawning. During the feeding period a positive energy balance was registered, an increase in total lipids was noted to 3.87% in September and to 4.63% in October, which is associated with intensive food consumption. According to the literature data, large-size atherina in the Middle Caspian Sea do not have empty intestines
in the spring-summer period, i.e. an increase in total muscle lipids in the autumn period is the result of fish nourishment earlier [4].
The statistical analysis revealed significant differences (p < 0.05) in the increase in the content of total lipids in the muscles of atherina in September, October in the feeding period relative to March by 29% and 41%, respectively, because in the spring period energy consumption for the formation and maturation of sexual products is carried out in the organism of atherina. In the autumn feeding period (October) there was a significant accumulation (p < 0.05) of total lipids compared to the pre-spawning and spawning stages (February-July), which is ensured by intensive food consumption.
With reference to total lipids, the amount of muscle water-soluble protein in atherina had a multidirectional character in different periods of the study.
In March in the pre-spawning period there was a slight increase in muscle protein to 58.40 mg/g compared to February (51.77 mg/g), which is explained by the predominance of protein synthesis over its decomposition. In April, with the beginning of spawning, the amount of water-soluble protein almost did not change and amounted to 58.17 mg/g. In July a decrease in protein in the muscle tissue of fish was revealed to 53.38 mg/g, due to its use as an energy substrate and removal from the body of atherina as part of sexual products. Similar metabolic mechanisms were noted in other fish with portioned spawning [17-19], which is
o
a. b
a
S' i
4
3
2
X X л
«
s с
explained by the redistribution of protein from the gonads to the muscles during the shedding of a repeated portion of the atherina roe. In September and October the amount of water-soluble protein does not increase, it slightly decreased to 49.51 and 46.94 mg/g, respectively, despite the consumption of food by atherina at these stages [4]. During the feeding period part of the protein in atherina is transformed into lipids, which accumulate intensively. During the studied periods of annual cycles there were no significant differences in the content of water-soluble protein in the muscle tissue of fish.
Thus, in the post-spawning - feeding period different directions of lipid and protein metabolism in the muscles were noted in atherina. Similar metabolic processes were registered at the feeding stage in Caspian sprat [18] and Azov anchovy [17].
In September a positive reliable (p < 0.05) statisti-
cal correlation was noted between the concentration of total muscle lipids and the mass of fish (r = +0.47), which characterized a positive energy balance in the organism necessary for the next stages of fish life.
Caspian atherina refers to fish species with portioned spawning, which causes a long duration of spawning and provides different possibilities to increase fertility. There exists some information in the literature about the fertility of atherina [20, 21], its high reliable correlation with the length and body weight of fish [4].
The reproduction of fish is based on the processes of development of the reproductive system - gameto-genesis. The reproductive system of atherina in the Caspian Sea is currently poorly studied.
The results of the study of fragments of the gonads of female atherina in February-March are shown in Fig. 2.
s о л
о
m
PQ
S
vo
S
s
Li
Fig. 2. Atherina oocytes at different stages of gonad maturation (SGM): a - SGM III-IV; b - SGM IV; c - resorption at SGM IV. Magnification 10 x 4
In February fish with completed SGM IV predominated (46%). Individuals in the amount of 34% had SGM III-IV and 20% of fish had SGM III-IV. At the same time, resorbing oocytes of the period of tropho-plasmic growth at an average level of 31% were noted in some species of atherina. The proportion of resorbing oocytes varied in the range of 17-47%.
In April the histological examination of fragments of the gonads of female atherina revealed that the largest number of fish (66%) were on SGM IV, individuals (20%) with gonads on SGM III-IV were noted. During this period atherina females (7%), that have started spawning, with gonads on SGM V, as well as individuals with ovaries on SGM VI-III (7%) were registered.
b
a
c
Disorders of the development of maturing roe in the range from 14 to 44% of oocytes with an average level of 30% were determined.
The individual analysis of female ovaries did not reveal mass resorption of oocytes. Thus, scattered ovarian resorption was recorded in atherina [22, 23], which was an adaptive mechanism of the organism [24, 25] allowing the fish to tolerate unfavorable conditions without losing the ability to reproduce.
According to literature data [26], fish with portioned spawning, if roe of the first portion was covered by mass resorption, have spawning, but it occurs already due to the second portion of roe and the loss of progeny can be in the range of 50-60%. In addition, unlike simultaneously spawning fish species, resorption does not have an inhibitory effect on the process of vitellogenesis in fish with portion spawning, i. e. two processes can simultaneously take place in the ovaries: generation and degeneration.
Therefore, the identified resorption processes in atherina are not pathological due to the fact that there is no effect on the development of other oocytes [23].
Conclusion
Therefore, morpho-physiological and biochemical studies of Caspian atherina were carried out at various stages of the annual cycle in the Caspian Sea.
According to the results of histological analysis, the beginning of spawning in atherina was registered in April. Mature oocytes with resorption at an average level of 30-31% were observed in the ovaries of portioned spawning atherina in February-April, which was the adaptive mechanism of the organism in this habitat.
In the pre-spawning period and at the beginning of spawning, changes in the quantitative composition of lipids and proteins characteristic of portioned spawning fish were noted aimed at spending the organism's energy for the formation, maturation of gonads, shedding of sexual products and ripening of the next portions of roe. At the stage of feeding (September-October) the amount of muscle water-soluble proteins and total lipids had a multidirectional character in the organism of atherina.
a*
o <
>
o
References
1. Khaldinova N. A. Materialy po razmnozheniiu i razvi-tiiu ryb v osolonennykh zalivakh Severnogo Kaspiia [Materials on the reproduction and development of fish in the saline bays of the Northern Caspian]. Trudy VNII morskogo rybnogo khoziaistva i okeanografii, 1951, vol. 18, pp. 99-126.
2. Berg L. S. Ryby presnykh vod SSSR i sopredel'nykh stran [Fresh water fish of the USSR and neighboring countries USSR]. Moscow, Izd-vo AN SSSR, 1949. Vol. 3. Pp. 1001-1004.
3. Taibov P. S., Kanieva N. A., Barabanov V. V., Ma-gomedova K. M. Promyslovo-biologicheskaia kharakteristika obyknovennoi kil'ki Clupeonella cultriventris caspia i ateriny Atherina mochon caspia v Tersko-Kaspiiskom rybokho-ziaistvennom podraione (Respublika Dagestan) v 2021 g. [Commercial and biological characteristics of the Caspian sprat Clupeonella cultriventris caspia and the atherina Atherina mochon caspia in the Tersk-Caspian Fisheries subdistrict (Republic of Dagestan) in 2021]. Biologicheskoe raznoobrazie: izuchenie, sokhranenie, vosstanovlenie, ratsional'noe ispol' zovanie: materialy III Mezhdunarodnoi nauchno-prakticheskoi konferentsii (Kerch', 13-18 sentiabria 2022 g.). Simferopol', OOO «Izdatel'stvo Tipografiia «Arial», 2022. Pp. 238-243.
4. Dzhabrailov Iu. M., Ustarbekova D. A,, Makhmudo-va N. N. Osobennosti biologii ateriny Atherina boyeri caspia (Eichwald, 1838) v zapadnoi chasti Srednego Kaspiia [Features of the biology of atherina Atherina boyeri caspia (Eichwald, 1838) in the western part of the Middle Caspian Sea]. Biora-znoobrazie flory i fauny Dagestana: materialy regional'noi nauchno-prakticheskoi konferentsii (Makhachkala, 20 marta 2012 g.). Makhachkala, Izd-vo DGPU, 2012. Pp. 108-110.
5. Taibov P. S., Ashumova S. G., Datsieva M. M., Medzhidova A. K. Morfoekologicheskaia kharakteristika ateriny Atherina boyeri Eichwald, 1838, v Tersko-Kaspiiskom rybokhoziaistvennom podraione [Morphoecological characteristics of atherina Atherina boyeri (Eichwald, 1838) in the Tersk-Caspian fishery subdistrict]. Kompleksnye issledovaniia biologicheskikh resursov iuzhnykh morei i rek: materialy IIIMezhdunarodnoi nauchno-prakticheskoi konferentsii mo-
lodykh uchenykh. Astrakhan', 2012. Pp. 111-113.
6. Dzhabrailov Iu. M., Ustarbekova D. A., Zurkhaeva U. D. Izmenchivost' morfologicheskikh priznakov ateriny Atherina boyeri caspia (Eichwald, 1838) [Variability of morphological signs of atherina Atherina boyeri caspia (Eichwald, 1838)]. Bioraznoobrazie flory i fauny Dagestana: materialy regional'noi nauchno-prakticheskoi konferentsii (Makhachkala, 20 marta 2012 g). Makhachkala, Izd-vo DGPU, 2012. Pp. 106-108.
7. Instruktsii po sboru i pervichnoi obrabotke materi-alov vodnykh bioresursov Kaspiiskogo basseina i sredy ikh obitaniia [Instructions for the collection and primary processing of materials of aquatic biological resources of the Caspian basin and their habitat]. Pod redaktsiei G. A. Suda-kova. Astrakhan', Izd-vo KaspNIRKh, 2011. 193 p.
8. Pravdin I. F. Rukovodstvo po izucheniiu ryb [Tutorial on the study of fish]. Moscow, Pishchevaia promyshlennost' Publ., 1966. 374 p.
9. Metody veterinarnoi klinicheskoi laboratornoi diag-nostiki: spravochnik [Methods of veterinary clinical laboratory diagnostics]. Pod redaktsiei prof. I. P. Kondrakhina. Moscow, KolosS Publ., 2004. 520 p.
10. Metody biologii razvitiia. Eksperimental'no-embriologicheskie, molekuliarno-biologicheskie i tsitolog-icheskie [Methods of developmental biology. Experimental-embryological, molecular-biological and cytological]. Pod redaktsiei T. A. Detlaf. Moscow, Nauka Publ., 1974. 619 p.
11. Prakticheskaia khimiia belka [Practical protein Chemistry]. Pod redaktsiei A. Darbre. Moscow, Mir Publ., 1989. 621 p.
12. Mikodina E. V., Sedova M. A., Chmilevskii D. A. Gistologiia dlia ikhtiologov. Opyt i sovety [Histology for ichthyologists. Experience and advice]. Moscow, Izd-vo VNIRO, 2009. 112 p.
13. Bavrina A. P. Sovremennye pravila primeneniia par-ametricheskikh i neparametricheskikh kriteriev v statistich-eskom analize mediko-biologicheskikh dannykh [Modern rules for the application of parametric and non-parametric
a. b
a
i ö i
ы
о Si
а
A
« §
о «
s с
ч о s
m
s -е-
о
•е р
о
m
Ш
m
<
у ü
criteria in the statistical analysis of biomedical data]. Vpomoshch' issledovateliu, 2021, no. 1 (66), pp. 64-73.
14. Bavrina A. P., Borisov I. B. Sovremennye pravila primeneniia korreliatsionnogo analiza [Modern rules for the application of correlation analysis]. V pomoshch' issledovateliu, 2021, no. 3 (68), pp. 70-79.
15. Kizevetter I. V. Biokhimiia syr'ia vodnogo pro-iskhozhdeniia [Biochemistry of raw materials of water origin]. Moscow, Pishchevaia promyshlennost' Publ., 1973. 424 p.
16. Maliarevskaia A. Ia. Obmen veshchestv u ryb v us-loviiakh antropogennogo evtrofirovaniia vodoemov [Metabolism in fish in conditions of anthropogenic eutrophica-tion of reservoirs]. Kiev, Naukova dumka Publ., 1979. 256 p.
17. Shul'man G. E. Fiziologo-biokhimicheskie oso-bennosti godovykh tsiklov ryb: monografiia [Physiological and biochemical features of annual cycles of fish]. Moscow, Pishchevaia promyshlennost' Publ., 1972. 368 p.
18. Kozlova N. V., Latunov A. A., Makarova E. G., Ma-khlun A. V. Fiziologo-biokhimicheskie izmeneniia kaspiiskoi obyknovennoi kil'ki (Clupeonella delicatula caspia Svetovidov) v razlichnye periody godovogo tsikla [Physiological and biochemical changes of the Caspian sprat (Clupeonella delicatula caspia Svetovidov) in different periods of the annual cycle]. Vestnik As-trakhanskogo gosudarstvennogo tekhnicheskogo universiteta. Seriia: Rybnoe khoziaistvo, 2021, no. 2, pp. 116-124.
19. Belianina T. N. Sezonnye izmeneniia zhirnosti belo-morskoi koriushki v sviazi s sozrevaniem gonad [Seasonal changes in the fat content of the White Sea smelt in connection with the maturation of the gonads]. Zakonomernosti chislennosti ryb Belogo moria. Moscow, Nauka Publ., 1966. Pp. 156-180.
20. Kazancheev E. N. Ryby Kaspiiskogo moria (opre-delitel') [Fish of the Caspian Sea (determinant)]. Moscow, Legkaia i pishchevaia promyshlennost' Publ., 1981. 167 p.
21. Iusufova Z. A. Znachenie ateriny Atherina mochon pontica natio caspia Eichwald v ispol'zovanii kormovoi bazy Kaspiiskogo moria. Avtoreferat dis. ... kand. biol. nauk [The
importance of atherina Atherina mochon pontica natio caspia Eichwald in the use of the feed base of the Caspian Sea. Ph.D. thesis]. Baku, 1969. 17 p.
22. Trusov V. E. Biologicheskaia kharakteristika i puti rybokhoziaistvennogo ispol'zovaniia osetrovykh, skapliva-iushchikhsia u Volgogradskoi plotiny [Biological characteristics and ways of fishery use of sturgeon accumulating at the Volgograd dam]. Osetrovye khoziaistva v vodoemakh SSSR. Moscow, Izd-vo AN SSSR, 1997. Pp. 143-150.
23. Kurdiaeva V. P., Shapavalov M. E. O rezorbtsii ootsi-tov i narusheniiakh v razvitii vosproizvoditel'noi sistemy u pred-stavitelei podsemeistva Sultrinae (semeistvo Ciprinidae) [On oocyte resorption and disorders in the development of the reproductive system in representatives of the subfamily Cultrinae (Family Ciprinidae)]. Izvestiia Tikhookeanskogo nauchno-issle-dovatel'skogo rybokhoziaistvennogo tsentra, 2002, vol. 131, pp. 390-408.
24. Gerbil'skii N. L. Ekologo-gistologicheskoe naprav-lenie v ikhtiologicheskikh issledovaniiakh [Ecological and histological direction in ichthyological research]. Trudy Vse-soiuznogo soveshchaniia po biologicheskim osnovam ry-bovodstva. Tomsk, 1959. Pp. 31-36.
25. Faleeva T. I. Analiz atrezii ootsitov u ryb v sviazi s adaptivnym znacheniem etogo iavleniia [Analysis of oocyte atresia in fish in connection with the adaptive significance of this phenomenon]. Voprosy ikhtiologii, 1965, vol. 5, iss. 3 (36), pp. 455-470.
26. Barkhalov R. M. Osobennosti razvitiia i funk-tsionirovaniia reproduktivnoi sistemy ryb (na primere se-meistva Cyprinidae) v izmenivshikhsia ekologicheskikh usloviiakh vodoemov Severo-Zapadnogo Kaspiia. Avtoreferat dis. ... kand. biol. nauk [Features of the development and functioning of the reproductive system of fish (on the example of the Cyprinidae family) in the changed ecological conditions of the reservoirs of the North-Western Caspian Sea. Ph.D. thesis]. Makhachkala, 2005. 26 p.
Список источников
1. Халдинова Н. А. Материалы по размножению и развитию рыб в осолоненных заливах Северного Каспия // Тр. ВНИИ мор. рыб. хоз-ва и океанографии. 1951. Т. 18. С. 99-126.
2. Берг Л. С. Рыбы пресных вод СССР и сопредельных стран. М.: Изд-во АН СССР, 1949. Т. 3. С. 1001-1004.
3. Таибов П. С., Каниева Н. А., Барабанов В. В., Маго-медова К. М. Промыслово-биологическая характеристика обыкновенной кильки Clupeonella cultriventris caspia и ате-рины Ather inamochon caspia в Терско-Каспийском рыбохо-зяйственном подрайоне (Республика Дагестан) в 2021 г. // Биологическое разнообразие: изучение, сохранение, восстановление, рациональное использование: материалы III Междунар. науч.-практ. конф. (Керчь, 13-18 сентября 2022 г.). Симферополь: ООО «Издательство Типография «Ариал», 2022. С. 238-243.
4. Джабраилов Ю. М., Устарбекова Д. А,, Махмудова Н. Н. Особенности биологии атерины Atherina boyeri caspia (Eichwald, 1838) в западной части Среднего Каспия // Биоразнообразие флоры и фауны Дагестана: материалы регион. науч.-практ. конф. (Махачкала, 20 марта 2012 г.). Махачкала: Изд-во ДГПУ, 2012. С. 108-110.
5. Таибов П. С., Ашумова С. Г., Дациева М. М., Меджидова А. К. Морфоэкологическая характеристика атерины Atherina boyeri Eichwald, 1838, в Терско-Каспийском рыбохозяйственном подрайоне // Комплекс-
ные исследования биологических ресурсов южных морей и рек: материалы III Междунар. науч.-практ. конф. молодых ученых. Астрахань, 2012. С. 111-113.
6. Джабраилов Ю. М., Устарбекова Д. А., Зурхаева У. Д. Изменчивость морфологических признаков атерины Atherina boyeri caspia (Eichwald, 1838) // Биоразнообразие флоры и фауны Дагестана: материалы регион. науч.-практ. конф. (Махачкала, 20 марта 2012 г.). Махачкала: Изд-во ДГПУ, 2012. С. 106-108.
7. Инструкции по сбору и первичной обработке материалов водных биоресурсов Каспийского бассейна и среды их обитания / под ред. Г. А. Судакова. Астрахань: Изд-во КаспНИРХ, 2011. 193 с.
8. Правдин И. Ф. Руководство по изучению рыб. М.: Пищ. пром-сть, 1966. 374 с.
9. Методы ветеринарной клинической лабораторной диагностики: справ. / под ред. проф. И. П. Кондрахина. M.: КолосС, 2004. 520 с.
10. Методы биологии развития. Экспериментально-эмбриологические, молекулярно-биологические и цитологические / под ред. Т. А. Детлаф. М.: Наука, 1974. 619 с.
11. Практическая химия белка / под ред. А. Дарбре. М.: Мир, 1989. 621 с.
12. Микодина Е. В., Седова М. А., Чмилевский Д. А. Гистология для ихтиологов. Опыт и советы. М.: Изд-во ВНИРО, 2009. 112 с.
13. Баврина А. П. Современные правила применения параметрических и непараметрических критериев в статистическом анализе медико-биологических данных // В помощь исследователю. 2021. № 1 (66). С. 64-73.
14. Баврина А. П., Борисов И. Б. Современные правила применения корреляционного анализа // В помощь исследователю. 2021. № 3 (68). С. 70-79.
15. Кизеветтер И. В. Биохимия сырья водного происхождения. М.: Пищ. пром-сть, 1973. 424 с.
16. Маляревская А. Я. Обмен веществ у рыб в условиях антропогенного эвтрофирования водоемов. Киев: Наукова думка, 1979. 256 с.
17. Шульман Г. Е. Физиолого-биохимические особенности годовых циклов рыб: моногр. М.: Пищ. пром-сть, 1972. 368 с.
18. Козлова Н. В., Латунов А. А., Макарова Е. Г., Махлун А. В. Физиолого-биохимические изменения каспийской обыкновенной кильки (Clupeonella delicatula caspia Svetovidov) в различные периоды годового цикла // Вестн. Астрахан. гос. техн. ун-та. Сер.: Рыбное хозяйство. 2021. № 2. С. 116-124.
19. Белянина Т. Н. Сезонные изменения жирности беломорской корюшки в связи с созреванием гонад // Закономерности численности рыб Белого моря. М.: Наука, 1966. С. 156-180.
20. Казанчеев Е. Н. Рыбы Каспийского моря (определитель). М.: Лег. и пищ. пром-ть, 1981. 167 с.
21. Юсуфова З. А. Значение атерины Atherina mochon pontica natio caspia Eichwald в использовании кормовой базы Каспийского моря: автореф. дис. ... канд. биол. наук. Баку, 1969. 17 с.
22. Трусов В. Е. Биологическая характеристика и пути рыбохозяйственного и спользования осетровых, скапливающихся у Волгоградской плотины // Осетровые хозяйства в водоемах СССР. М.: Изд-во АН СССР, 1997. С. 143-150.
23. Курдяева В. П., Шапавалов М. Е. О резорбции оо-цитов и нарушениях в развитии воспроизводительной системы у представителей подсемейства СиИгтае (семейство Ciprinidae) // Изв. Тихоокеан. науч.-исследоват. ры-бохозяйств. центра. 2002. Т. 131. С. 390-408.
24. Гербильский Н. Л. Эколого-гистологическое направление в ихтиологических исследованиях // Тр. Все-союз. совещ. по биолог. основам рыбоводства. Томск, 1959. С. 31-36.
25. Фалеева Т. И. Анализ атрезии ооцитов у рыб в связи с адаптивным значением этого явления // Вопр. ихтиологии. 1965. Т. 5. Вып. 3 (36). С. 455-470.
26. Бархалов Р. М. Особенности развития и функционирования репродуктивной системы рыб (на примере семейства Cyprinidae) в изменившихся экологических условиях водоемов Северо-Западного Каспия: автореф. дис. ... канд. биол. наук. Махачкала, 2005. 26 с.
н i
A
<i V
The article was submitted 12.04.2023; approved after reviewing 31.05.2023; accepted for publication 31.08.2023 Статья поступила в редакцию 12.04.2023; одобрена после рецензирования 31.05.2023; принята к публикации 31.08.2023
Information about the authors / Информация об авторах
Pirmurad S. Taibov - Head of the Sector of Commercial Ichthyology; West-Caspian Department of the Volga-Caspian Branch Russian Federal "Research Institute of Fisheries and Oceanography"; taibov.p@yandex.ru
Пирмурад Султанмурадович Таибов - заведующий сектором промысловой ихтиологии; Западно-Каспийский отдел Волжско-Каспийского филиала Всероссийского научно-исследовательского института рыбного хозяйства и океанографии; taibov.p@yandex.ru
о О
Nuria A. Kanieva — Doctor of Biological Sciences, Professor; Professor of the Department of Applied Biology and Microbiology; Astrakhan State Technical University; kanievana52@mail.ru
Vitali V. Barabanov — Candidate of Biological Sciences; Head of the Resource Research Center; Volga-Caspian Branch Russian Federal "Research Institute of Fisheries and Oceanography"; barabanov2411@yandex.ru
Нурия Абдрахимовна Каниева - доктор биологических наук, профессор; профессор кафедры прикладной биологии и микробиологии; Астраханский государственный технический университет; kanievana52@mail.ru
Виталий Викторович Барабанов - кандидат биологических наук; руководитель Центра ресурсных исследований; Волжско-Каспийский филиал Всероссийского научно-исследовательского института рыбного хозяйства и океанографии; barabanov2411@yandex.ru
i R
Natalia V. Kozlova - Candidate of Biological Sciences; Head of the Laboratory of Molecular Genetics and Physiology; Volga-Caspian Branch Russian Federal "Research Institute of Fisheries and Oceanography"; natali19_12@mail.ru
Наталья Викторовна Козлова - кандидат биологических наук; заведующий лабораторией молекулярной генетики и физиологии; Волжско-Каспийский филиал Всероссийского научно-исследовательского института рыбного хозяйства и океанографии; natali19_12@mail.ru
