ИЗМЕНЕНИЕ СТРУКТУРЫ ПОПУЛЯЦИИ ЖАБРОНОГОГО РАКА ARTEMIA SP. В АКТАШСКОМ ОЗЕРЕ (КРЫМСКИЙ ПОЛУОСТРОВ) В ОСЕННЕ-ЗИМНИЙ ПЕРИОД (ПО ДАННЫМ 2018 Г.)
Авторы:
1.
Керченский государственный морской технологический университет
(кандидат биологических наук; заведующий кафедрой водных биоресурсов и марикультуры)
2.
Керченский государственный морской технологический университет
(лаборант кафедры водных биоресурсов и марикультуры)
3.
Азово-Черноморский филиал Всероссийского научно-исследовательского института рыбного хозяйства и океанографии
(кандидат биологических наук; заведующий лабораторией биоресурсов внутренних водных объектов)
Россия
Россия
4.
Керченский государственный морской технологический университет
(кандидат биологических наук; доцент кафедры водных биоресурсов и марикультуры)
Россия
Россия
5.
Карадагская научная станция им. Т. И. Вяземского, природный заповедник РАН, филиал федерального исследовательского центра «Институт биологии южных морей им. А. О. Ковалевского РАН»; Керченский государственный морской технологический университет
(кандидат биологических наук; старший научный сотрудник лаборатории биохимии и физиологии гидробионтов;доцент кафедры водных биоресурсов и марикультуры)
Россия
Россия
Тип:
Статья
Страницы:
с 16
по 28
Статус:
Опубликован
Получено:
20.09.2022
Одобрено:
20.09.2022
Опубликовано:
28.09.2022
Классификаторы:
УДК 595.3 Branchiata (Crustacea). Ракообразные
Язык материала:
английский
Ключевые слова:
жаброногие раки, Artemia, Акташское озеро, структура популяции, осенне-зимний период, цисты, условия среды
Аннотация (русский):
Представлены результаты исследования изменения возрастной (по всем онтогенетическим группам) и половой структуры, а также продуцирования цист в двух популяциях Artemia sp., обитающих в различных контрастно-различающихся (по площади водного объекта, его глубине, солености воды, характеру донных отложений, уровню антропогенной нагрузки) условиях среды гипергалинных водоемов аквального комплекса Акташский. Приведены данные о динамике размерного состава каждой из половозрастных групп (науплиусов, метанауплиусов, ювенильных и предвзрослых раков, самок с овисаками и без них, самцов). Выполнены измерения цист (Dmax и Dmin). Особенностью данной работы является время проведения наблюдений (осень-начало зимы) – период «угасания» популяции. На примере двух изолированных водоемов исследовалась взаимосвязь изменений в популяциях в условиях снижения показателей наиболее значимых абиотических факторов среды – солености воды и ее температуры. Получены данные об условиях продуцирования цист для водоемов данной группы. Продукция жаброногих раков (г/м3) в водоемах не зависит от объема в них воды. Популяции Artemia в исследуемых водоемах имееют широкий диапазон зоны оптимума по солености среды. Установлено, что главным пусковым показателем начала зимней паузы жизнедеятельности популяции является температура воды (8–10 °С), при этом соленость имеет меньшее значение. Приведены данные о составе и численности прочих видов беспозвоночных животных (личинок двукрылых, имаго жесткокрылых, копепод), обитающих в исследуемых водных объектах в меженный осенне-зимний период.
Представлены результаты исследования изменения возрастной (по всем онтогенетическим группам) и половой структуры, а также продуцирования цист в двух популяциях Artemia sp., обитающих в различных контрастно-различающихся (по площади водного объекта, его глубине, солености воды, характеру донных отложений, уровню антропогенной нагрузки) условиях среды гипергалинных водоемов аквального комплекса Акташский. Приведены данные о динамике размерного состава каждой из половозрастных групп (науплиусов, метанауплиусов, ювенильных и предвзрослых раков, самок с овисаками и без них, самцов). Выполнены измерения цист (Dmax и Dmin). Особенностью данной работы является время проведения наблюдений (осень-начало зимы) – период «угасания» популяции. На примере двух изолированных водоемов исследовалась взаимосвязь изменений в популяциях в условиях снижения показателей наиболее значимых абиотических факторов среды – солености воды и ее температуры. Получены данные об условиях продуцирования цист для водоемов данной группы. Продукция жаброногих раков (г/м3) в водоемах не зависит от объема в них воды. Популяции Artemia в исследуемых водоемах имееют широкий диапазон зоны оптимума по солености среды. Установлено, что главным пусковым показателем начала зимней паузы жизнедеятельности популяции является температура воды (8–10 °С), при этом соленость имеет меньшее значение. Приведены данные о составе и численности прочих видов беспозвоночных животных (личинок двукрылых, имаго жесткокрылых, копепод), обитающих в исследуемых водных объектах в меженный осенне-зимний период.
Ключевые слова:
жаброногие раки, Artemia, Акташское озеро, структура популяции, осенне-зимний период, цисты, условия среды
жаброногие раки, Artemia, Акташское озеро, структура популяции, осенне-зимний период, цисты, условия среды
Introduction
Branchiopods Artemia Leach, 1819 (Crustacea: Branchiopoda: Anostraca), or brine shrimp, are inhabitants of continental hyperhaline water bodies. Inhabiting water bodies with sometimes extreme living conditions, having an incompletely elucidated species composition of the genus, possessing a complex ontogenetic development cycle, amazing viability of dormant eggs (cysts), as well as alternating sexual and parthenogenetic ways of reproduction, this group of living creatures has always attracted the attention of biologists [1-18]. The increased interest in crustaceans Artemia is also emphasized by the fact that their cysts are of commercial importance, being a strategically im-portant starting (primary) fodder for juveniles of most fish species rearing in aquaculture. The ever-increasing economic need for the brine shrimp cysts is realized exclusively by their removal (catch) from natural populations [16-19]. Dynamically changing climatic conditions lead to sharp fluctuations in the number of brine shrimp in water bodies throughout the season, contributing to a certain unpredictability in forecasting and providing of its catch. At the same time, attempts to cultivate brine shrimp in artificial conditions in order to produce their cysts on an industrial scale have not yet overcome the scope of laboratory experiments. In this regard, the study of the nature of changes in population processes in natural water bodies, as well as their relationship with the main abiotic and biotic processes, the results of which make it possible to justify the rational use of Artemia sp. stocks, do not lose their relevance today.
Spreading of crustaceans Artemia in the water bodies of Crimea is extremely uneven, both geographically in terms of the location of hyperhaline water bodies and in time. Often water bodies in which these crustaceans live mostly have extremely shallow depths due to the seasonal distribution of precipitation. They quickly desalinate during heavy rains or, conversely, dry up during long dry periods, which leads to the ces-sation of the development of brine shrimp and its disappearance [1, 2, 8-11].
In the eastern part of the Crimean Peninsula Artemia sp., or brine shrimp, lives in almost all water bodies with permanent or seasonal high water salinity with values from 40 to 300‰. For some water bodies, with contrasting hydrological characteristics changing throughout the year, the development of Artemia sp. is characterized by short-term outbreaks of population development. For other water bodies with relatively constant conditions the long period of populations existence can take place.
Lake Aktashskoe is located in the northern part
of the Kerch Peninsula (Crimea). As a result of its economic use, it is divided into a number of separate reservoirs with contrastingly different morphological characteristics (area, depths, nature of the banks and bed, interconnection with other water bodies) and hydrological regime (provision of surface runoff, water level dynamics) [20]. So, having the complicated structure, this complex is a typical hyperhaline reservoir and is most suitable for performing model population studies.
The object of our study was the population of the branchiopod crustacean Artemia sp. inhabiting the reservoirs of the hyperhaline aquatic complex – Lake Aktashskoe. And the subject of the study is the structure of the population of this crustacean and its relationship with abiotic environmental factors.
The novelty of this research lies in the fact that, despite the seemingly high degree of study of the brine shrimp, there can be found very poor data on the state of the populations of branchiopod crustacean in the lakes of the Crimea in the autumn-winter period, and there is no data at all on the complex of hyperhaline reservoirs of the Kerch Peninsula. Studies of the brine shrimp in the framework of the rational use of its commercial stocks are limited only to the period
of late spring – early autumn.
The aim of this work is to study the population structure of the branchiopod crustacean Artemia sp. in non-drying water bodies of the hyperhaline aquatic complex of Lake Aktashskoe in the autumn-winter period of 2018/2019.
To achieve this aim, the following tasks were formulated:
1. To determine control stations (typical parts
of the reservoir that are contrasting in terms of abiotic conditions) and carry out sampling of the brine shrimp in the autumn-winter period, as well as to collect data on the habitat environment (salinity, temperature) in the reservoir.
2. To perform cameral processing of collected samples of the brine shrimp to determine their qualitative (population structure by age groups and sex) and quantitative (number and biomass of each of the groups) structure.
3. To analyze the relationship of changes in the structure of the brine shrimp population (by age and sex) under the influence of the key abiotic environ-mental factors (salinity and water temperature) occurring in the autumn-winter period. To study the size structure of the Artemia sp. population and its changes in the second half of the vegetation season.
4. To determine the composition of other species
of invertebrates living in the control hypersaline reservoirs of Lake Aktashskoe complex.
Materials and methods
The material for the study was seven series of zooplankton samples from two water bodies of Lake Aktashskoe complex (Fig. 1), sampled in September-December 2018.
Fig. 1. Sampling sites at Lake Aktashskoe: 1 – cooling pond; 2 – natural part
The first pond (No. 1 – the cooler) is a hydrotechnical structure lined with an artificial dam, intended for cooling heat-exchange waters of the Crimean nuclear power plant designed at the end of the 20th century (construction was not finished). It is characterized by a significant water surface area (7-9 km2) and comparatively big depth (2.0 m). Its bottom is relatively hard and clayish. The second pond (No. 2 – the natural part) is a part of the lake practically untouched by human activity. It is of much smaller area (up to 1.5 km2) and average depth (0.7 m). Bottom of this water body is covered with thick layer of silt.
The control water bodies practically do not have their own catchment area, their water supply occurs mainly with the groundwater and, to a much lesser extent, from atmospheric precipitation. Pond No. 2 has an intermittent connection with the waters of the Kazantip Bay (Sea of Azov) through an artificial channel. Despite significant fluctuations in the water level in both reservoirs (during the time of observations of the authors in No. 1-0.7 m; in No. 2-0.5 m), they do not dry out throughout the year, and therefore it is possible to study the entire annual cycle of development of the Artemia sp. population.
Basic characteristics of water environment in the ponds when zooplankton was sampled are shown in Table 1.
Table 1
Water temperature and salinity in the ponds when sampling zooplankton
Pond Parameters Dates of zooplankton sampling
30.09.2018 15.10.2018 18.11.2018 15.12.2018
No. 1 Temperature, °С 19.0 16.0 7.8 3.0
Salinity, ‰ 204 196-200 208 186
No. 2 Temperature, °С 26.0 – 9.4 4.0
Salinity, ‰ 140 – 112 76
Sampling was carried out using the Plankton Net acc. to Apstein by filtering 100 liters of water from the water body, taken along a transect with a length of at least 100 m. The samples taken were fixed immediately with a 4% formaldehyde solution. The water temperature (with a mercury thermometer) and its salinity (with a Kelilong RHS-28ATC refractometer) were determined at the same time.
Sample processing was carried out in the laboratory according to the generally used methods. Quantitative accounting of different sex and age groups
of brine shrimp was carried out using a Bogorov camera, the determination of the individual weight of brine shrimp was carried out using reference tables of the Russian Federal Research Institute of Fisheries and Oceanigraphy [19].
The size of the brine shrimp was determined by measurements of at least 20 specimens of each of its size and age groups in each of the samples in the divisions of the eyepiece micrometer. The actual dimensions in micrometers were calculated after calibrating the division value against the micrometer object.
Statistical processing of the obtained data was carried out using the Microsoft Excel 2010 and IBM SPSS Statistics v. 22 software packages.
Results
Status of the Artemia sp. population in the cool-ing pond (No. 1) of Lake Aktashskoe. The Artemia sp. population in the part of Lake Aktashskoe that is transformed into a cooling reservoir is of a complex structure, as it is proved by the data on number and biomass of its different sex and age groups (Table 2).
Table 2
Change in the number and biomass of the Artemia sp. by age groups
in the cooling pond (No. 1) during September-December 2018
Ages groups, gender Number, ind./m3 Biomass, g/m3
30.09.2018 15.10.2018 18.11.2018 15.12.2018 30.09.2018 15.10.2018 18.11.2018 15.12.2018
Cysts 1 053 6 695 4 455 205 922 0.01 0.07 0.04 2.06
Nauplii 21 32 560 0.00 0.00 0.00 0.01 0.00
Metanauplii 80 858 1 643 0.00 0.01 0.13 0.25 0.00
Juvenile 210 242 47 0.00 0.11 0.13 0.03 0.00
Pre-adult 1 941 137 2 699 553 3.32 0.23 4.62 0.95
Females without ovi-sacs 312 137 5 9 0.80 0.35 0.01 0.02
Females with ovisacs 244 447 68 12 0.90 1.65 0.25 0.04
Females 556 584 73 21 1.70 2.00 0.26 0.06
Males 240 258 1 251 141 0.66 0.71 3.45 0.39
Total 4 101 8 801 10 732 206 637 5.81 3.27 8.66 3.46
In the samples, except the sample taken in December, all age stages are recorded. The number of individuals per 1 m3 in groups is different, according to the dates of sampling.
In September-October under conditions of a gradual but insignificant temperature decrease and a relatively stable salinity within 200‰, the number of younger (except for metanauplii) and older age groups changed little (Fig. 2). In early November there was a rapid increase of brine shrimp number in general – the proportion of younger groups increases many times
(Table 2, Fig. 2).
Fig. 2. Change in age structure of the Artemia sp. population in the cooling reservoir (No. 1)
In November, there is a sharp (more than 2 times) decrease in temperature, which affects the structure
of the branchiopod population. In the November plankton a huge amount of dead Artemia sp. is recorded, especially in older age groups. Subsequently, juveniles also die in the mass, and the number of brine shrimp in December goes to zero.
An analysis of the data on sex structure of the Artemia sp. population is shown at the Fig. 3.
Fig. 3. Change in sex structure of the Artemia sp. population from the cooling pond (No. 1)
Their ratio also changed when the main abiotic factors (temperature, salinity) changed during the study period. At the same time, the proportion of females, including those with ovisacs, as well as males, their ratio, may indicate a change (predominance) of parthenogenetic or sexual reproduction in the population.
During September parthenogenetic reproduction prevails in the brine shrimp population at the water body
No. 1. The presence of cysts in plankton indicates the existence of a sexual mode of reproduction in this period. The ratio of females and males in September and October is stable, at the level of 2 : 1 (Fig. 3). At the same time, females with ovisacs make up about half of their number in September, but in October the part of females with ovisacs is already more than 75% (Fig. 3).
With abrupt decrease in temperature in the water body in November, the proportion of adults is significantly reduced by less than 10% of their earlier number. But at the same time, the ratio of females and males changes vice versa amounting to 1 : 17. Perhaps, this fact indicates an increasing role of sexual reproduction. Parthenogenetic females die, and their place is taken by growing females that reproduce bisexually. This is also confirmed by the increasing number of the brine shrimp cysts in plankton. It is important that females predominated in the mass of dead mature individuals in November, and proportion of males was disproportionately smaller. In December the proportion of females slightly increases.
The number and biomass of the Artemia sp. cysts in the cooling pond (No. 1) during the autumn-winter period changes significantly (Fig. 4).
Fig. 4. Change in the number and biomass of the Artemia sp. cysts in the cooling pond (No. 1)
In September, the smallest number of cysts was recorded (1 053 cysts/m3), in October, as the water temperature decreases, the proportion of cysts increases (6 695 cysts/m3), and in November it again slightly decreases (4 455 cysts/m3). This may be a consequence of the death of most of the females, including those bred bisexually. The maximum number of cysts was recorded in December 2018 (205 922 cysts/m3). Thus, during the study, the biomass of brine shrimp cysts from September to December increased 204 times and amounted to 2.04 g/m3.
According to the data obtained from the reservoir No. 1 (part of Lake Aktashskoe), it should be noted that the starting point (limiting condition) for the beginning of changes in the structure of the population (by age groups and sex), preceding the winter pause, is a decrease in water temperature up to 10°C and lower at relatively stable salinity values (at the level
of 200‰). Under these conditions, Artemia begins an increased production of dormant eggs – cysts.
Status of the Artemia sp. population at the natural part of Lake Aktashskoe (pond No. 2)
The brine shrimp population at the natural, no transformed part of Lake Aktashskoe (pond No. 2) also is of a complex age structure represented by all groups. However, their one-time ratio and quantitative indicators are different (Table 3).
Table 3
Change in the number and biomass of the Artemia sp. by age groups during September-December 2018
at the natural part of Lake Aktashskoe (pond No. 2)
Ages groups, gender Number, ind./m3 Biomass, g/m3
30.09.2018 18.11.2018 15.12.2018 30.09.2018 18.11.2018 15.12.2018
Cysts 85 404 94 823 1 068 263 0.85 0.95 10.68
Nauplii 22 20 32 0.00 0.00 0.00
Metanauplii 4 730 20 64 0.71 0.00 0.01
Juvenile 19 004 109 127 10.45 0.06 0.07
Pre-adult 3 860 40 0.00 6.60 0.07 0.00
Females without ovisacs 1 404 385 158 3.59 0.99 0.40
Females with ovisacs 4 969 1 283 600 18.38 4.75 2.22
Females 6 373 1 668 758 21.97 5.74 2.62
Males 183 356 63 0.50 0.98 0.17
Total 119 576 97 036 1 069 307 41.08 7.80 13.55
The largest number of all groups (except cysts) was registered in September at water temperature of 26°C
and salinity of 140‰ (Fig. 5).
Fig. 5. Change in the age structure of the Artemia sp. population at the natural part of Lake Aktashskoe (pond No. 2)
Subsequently, under lower temperatures and desalination (decrease in salinity) of water, the number
of each of the groups decreased significantly (by 9 or more times).
The structure of the brine shrimp population by sex in the natural part of the lake has significant imbalance towards females (Fig. 6).
Fig. 6. Change in the sex structure of the Artemia sp. population at the natural part of Lake Aktashskoe (pond No. 2)
The ratio of females and males over the control periods varies from 5 : 1 to 34 : 1 with an obvious tendency to increase the proportion of males in the second half of the observation period (November-December 2018). The part of females with ovisacs throughout the entire period is consistently high – 77-79%.
According to the structure of the population by sex, it is possible to assume that Artemia sp. at the pond No. 2 during the entire period of observations reproduces mainly parthenogenetically. Although the number of cysts is quite high. That is, there are two groups of females in the water body, using different methods of reproduction (reproductive strategies).
Cyst production in the brine shrimp population
of the natural part of Lake Aktashskoe remains practically unchanged under conditions of salinity decrease from 140 to 112‰, and water temperature from 26 to 9°C (Fig. 7).
At salinity values below 110‰ and temperatures of 8°C, the number of cysts in plankton rapidly increases.
Fig. 7. Change in the number and biomass of the Artemia sp. cysts in the natural part (No. 2) of Lake Aktashskoe
Analysis of the size composition of the population in various water bodies of the aquatic complex – Lake Aktashskoe
In processing of samples there were measured the
body length of brine shrimp in each of the sex and age groups, as well as the sizes (Dmax, Dmin, S) of its cysts. The measurement results are shown in Table 4.
When comparing the size of brine shrimp at certain stages of development in ponds 1 and 2 (Fig. 8), according to the dates of selection, at first glance, a chaotic dynamic is observed.
a
b
Fig. 8. Change in the average length of the Artemia sp. by groups:
a – in the cooling pond; b – in the natural part of Lake Aktashskoe
However, it is possible to identify a number of specific trends. The younger age groups (nauplius – metanauplius, and at pond 2, juveniles) become smaller. Artemia sp. during the pre-adult stage grows faster up to the end of the season. In the cooling pond (No. 1) adult females get smaller in size when temperature decreases, while in the natural part of the lake (pond No. 2), on the contrary, they become larger. The sizes of males in both ponds practically do not change throughout the entire observation period (without taking into account the values for November in pond No. 1).
In general, the size of the Artemia sp. in terms
of stages of development in both ponds is relatively equal. The only thing that requires explanation is the fact of a sharp difference in the size of males in reservoirs No. 1 (4.73-5.25 mm) and No. 2 (6.10-6.44 mm). The issue of changes in the size of females during the period of deterioration of the living conditions of the population remains debatable. What is its cause: accelerates (or slows down) their differentiation by sex and their participation in reproduction; slows down (or accelerates) their growth (food supply, etc.)?
When comparing the average values for groups in September (the period of relative well-being of the population) and November (the beginning of mass production of cysts and depression of the population) for ponds (Fig. 9), the following was established.
a b
Fig. 9. Change in the average size of the brine shrimp in groups as it grows in September
and November 2018: a – at the cooling pond; b – at the natural part of Lake Aktashskoe
In the cooling pond (No. 1) under conditions of
a sharp temperature drop, the length of the Artemia sp. specimens decreases in all groups. In the natural part (No. 2), on the contrary, there are no significant changes in the values of the average body length in any of the groups.
Other species of invertebrates in hyperhaline ponds of the aquatic complex – Lake Aktashskoe
In the samples of zooplankton from the cooling pond (No. 1) at a salinity of 186-200‰, in addition to brine shrimp, a few individuals of one species of copepods were found: October 15, 2018 – 5 ind./m3; December 15, 2018 – 3 ind./m3.
In the natural part (No. 2) of Lake Aktashskoe the biological diversity of other invertebrate species was significantly higher. Seven species of invertebrates were recorded here, including 5 species of larvae
of near-aquatic insects, one species of beetles (imago) and 1 species of copepods. The numbers in species are given in Table 5.
Table 5
Composition and dynamics of other species of invertebrates in the plankton of the natural part (No. 2)
of Lake Aktashskoe during the study period
Aquatic arthropods,
juveniles and adults Number, ind./m3
30.09.2018 (salinity 140‰) 18.11.2018 (salinity 112‰) 15.12.2018 (salinity 76‰)
Chironomidae larvae 611 119 95
Larvae of other Diptera 7 0 32
Coleoptera 14 0 0
Copepoda 197 375 2 874
In addition, in September-November, the presence of green filamentous algae was noted in the plankton of the pond (No. 2), while their number decreased from the beginning of the study period to its end (there were no algae in December).
Conclusion
Sex and age structure of populations of the bran-chiopod crustacean Artemia sp. in the autumn-winter period, it has a complex structure, which is due both to environmental conditions and too much greater adaptive abilities of this species than is commonly believed. The nature of the impact on brine shrimp of the accepted triggering factors of the natural environment (threshold values of water temperature and salinity), which predetermine the decline in population size during the transition from parthenogenetic reproduction
of offsprings to sexual reproduction or, on the contrary, rapid increase of brine shrimp caused by the release of nauplii from wintering cysts, does not can always be unambiguously used in practice.
In both ponds (No. 1 – cooling pond, No. 2 – natural part of Lake Aktashskoe), at different salinity at water temperatures below 10-8°C, an increase in the number of cysts takes place. At the same time, the difference in salinity in water bodies is almost 100‰. Apparently, this indicator in brine shrimp has a much wider range of optimum.
The Artemia sp. population in the cooling pond
(No. 1), due to its larger volume of water and depth, develops gradually, without sharp changes in numbers and short-term disappearance of certain age groups.
At the same time, the productivity of this population is somewhat lower than in the second pond, which is probably due to increased salinity and food shortage, etc. On the contrary, the population in the natural part (No. 2) of the lake, which has a smaller volume and depth, develops explosively, but the biomass of the Artemia sp. at the peak of its development is much higher. Under conditions of a sharp decrease in temperature, the nature of changes in the sex, age and size structure in water bodies with different salinity is different.
The nature of regulation of the Artemia sp. population, its sex and size-age structure are probably much more complicated than it was previously thought. At the same time, it is necessary to take into account a larger list of environmental factors, as well as the nature
of their joint influence, which requires additional research involving more analytical material in terms
of chronology (duration of observation) and geography.
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