Morphological and genetic variation of Artemia sinica in Yuncheng Salt Lake over the past 30 years
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摘要: 卤虫是海水鱼类及甲壳类苗种培育中重要的活饵料，90%以上来源于野生卤虫资源。为探究近30年环境变化与独特养殖管理模式对中国运城盐湖卤虫形态学及遗传特征的影响，本研究对1993年（YC-1993）、2019年（YC-2019）和2023年（YC-2023）采自运城盐湖的卤虫卵进行孵化并饲养至成体，进行卤虫卵和成虫的形态学测量；同时采用特定位点扩增片段测序（SLAF-seq）技术进行群体遗传学分析。结果表明，YC-2023群体的体长、腹长、卵囊宽、复眼间距、复眼直径、第二触角长、抱握器外周长等形态学指标上均极显著大于YC-1993群体，而YC-2023群体卵径显著小于YC-1993群体。遗传分析显示，YC-2023群体的遗传多样性最低；三个群体的多态信息含量（PIC）介于0.252−0.305之间，属于中度多态（0.25<PIC<0.5）。YC-2019和YC-2023群体间的遗传分化系数Fst值为0.087，属于中度遗传分化（0.05<Fst<0.15）。而YC-1993与另外两个种群间的Fst值为0.151，属于高度遗传分化（0.15<Fst<0.25）。系统发育树、主成分分析、亲缘关系热图及Admixture分析进一步揭示，虽然三个群体间存在一定遗传分化，但均起源于单一共同祖先。运城卤虫的形态性状与遗传变异，可能源于运城盐湖近年来经历的环境变化以及独特养殖和管理模式所导致的遗传选择与遗传漂变。本研究为运城盐湖卤虫种质资源的保护与利用提供了理论支撑。Abstract: Artemia is important live bait in the cultivation of marine fish and crustacean seedlings, with more than 90% derived from wild Artemia resources. In order to study the effects of environmental changes and the unique aquaculture and harvest management on the morphometrical and genetic characteristics of Artemia in Yuncheng Salt Lake, Shanxi Province in the past 30 years, Artemia cysts collected in 1993 (YC-1993), 2019 (YC-2019) and 2023 (YC-2023) from Yuncheng Salt Lake were cultured and morphometrical measurements were conducted. Specific-locus amplified fragments sequencing (SLAF-seq) was used for population genetics analysis. The results showed that the YC-2023 group was extremely significantly larger than the YC-1993 group in terms of body length, abdominal length, ovary width, interocular distance, eye diameter, second antenna length, and peripheral claspers, while the cyst diameter of the YC-2023 group was significantly smaller than that of the YC-1993 group. The results of genetic analysis showed that the YC-2023 group had the lowest genetic diversity. The polymorphism information content (PIC) of three populations was between 0.252 and 0.305, showing moderate polymorphism (0.25<PIC<0.5). The Fst value of the genetic differentiation coefficient between the YC-2019 and YC-2023 populations is 0.087, indicating moderate genetic differentiation (0.05<Fst<0.15). However, the Fst value between the YC-1993 and the other two populations is 0.151, indicating a high degree of genetic differentiation (0.15<Fst<0.25). Phylogenetic tree, principal component analysis, the kinship heat map and Admixture analysis further revealed that although there was a certain degree of genetic differentiation among the three populations, their genetic information originated from the same original ancestor. The morphometrical traits and genetic variations of Yuncheng Artemia may result from genetic selection and genetic drift caused by the environmental changes that Yuncheng Salt Lake has undergone in recent years, and unique aquaculture and harvest management. This study will provide theoretical support for the conservation and utilization of Artemia germplasm resources in Yuncheng Salt Lake.
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Key words:
- Yuncheng Salt Lake /
- Artemia /
- genetic diversity /
- genetic structure /
- germplasm
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图 1 三个卤虫群体的形态学差异分析
F. 雌性; M. 雄性.*表示两组间差异显著（P < 0.05），**表示两组间差异非常显著（P < 0.01），***表示两组间差异极显著（P < 0.001）
Fig. 1 Morphometrical variation analysis of three Artemia populations
F. Female, M. Male. *indicates a significant difference between the two groups (P < 0.05), ** indicates a very significant difference between the two groups (P < 0.01), and *** indicates an extremely significant difference between the two groups (P < 0.001)
图 2 三个卤虫群体形态学参数主成分分析
A. 雌性; B. 雄性; a. 体长, b. 腹长, c. 体宽, d. 复眼间距, e. 复眼直径, f. 头宽, g. 生殖节, h. 抱握器外周长, i. 额结节, j. 卵囊宽, k. 第二触角长.
Fig. 2 Principal component analysis of morphometrical data of three Artemia populations
A. Female; B. Male; a. Body length, b. Abdominal length, c. Body width, d. Distance between compound eyes, e. Diameter of compound eyes, f. Head width, g. Genital segment, h. Outer circumference of the clasping organ, i. Frontal tubercle, j. Ovary width, k. Length of the second antenna.
图 3 基于SNP的三个卤虫群体主成分分析（A）；三个卤虫群体的邻接系统发育树（B）
Fig. 3 (A) Principal component analysis of three Artemia populations based on SNPs; (B) Neighbor-Joining phylogenetic tree of three Artemia populations
图 4 Admixture 分析的群体分组结果（K=1 至 10）（A）；交叉验证错误率（B）
Fig. 4 (A) Model-based population assignment by Admixture analysis for K=1 to 10; (B) Cross validation error rate
图 5 三个卤虫群体的亲缘关系热图
图中颜色梯度对应亲缘系数，颜色由蓝色向红色过渡表示亲缘关系系数逐渐升高，个体间遗传相似度逐渐增强。
Fig. 5 Heat map of kinship of three Artemia populations
The color gradient in the figure corresponds to the coefficient of kinship. The transition from blue to red indicates a gradual increase in the coefficient of kinship, meaning the genetic similarity between individuals gradually strengthens.
表 1 样品来源及相关信息
Tab. 1 Sample source and information
缩写 样品来源 采集年份 AR-ARC种质库编号 物种名称 YC-1993 中国运城盐湖 1993 CHN-SL-29 A. sinica YC-2019 中国运城盐湖 2019 CHN-SL-66 A. sinica YC-2023 中国运城盐湖 2023 CHN-SL-94 A. sinica 
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表 2 三个卤虫群体的遗传多样性指数
Tab. 2 Genetic diversity indexes of three Artemia populations
群体 期望等位基因数（Ea） 观测等位基因（Oa） 期望杂合度（He） 观测杂合度（Ho） 基因多样性指数（Nei） 多态性信息含量（PIC） 香农-威纳指数（I） YC-1993 1.554 1.850 0.374 0.421 0.409 0.297 0.553 YC-2019 1.519 1.769 0.385 0.286 0.423 0.305 0.567 YC-2023 1.357 1.700 0.310 0.375 0.310 0.252 0.476
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