东海剑尖枪乌贼耳石微结构及生长特性研究
Study on microstructure and growth characteristics of Uroteuthis edulis statolith in East China Sea
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摘要: 文章根据2017年和2018年9月至翌年3月在东海采集的剑尖枪乌贼 (Uroteuthis edulis) 样本，对其耳石形态进行观察与测量，并结合日龄信息对耳石微结构及生长特征进行了研究。结果表明，耳石生长纹由明暗相间的环纹组成，依据生长纹的宽度及亮度划分为后核心区、暗区和外围区。通过耳石日龄进行逆推发现春、夏产卵群体为优势群体，优势群体间耳石各形态参数差异显著 (P<0.05)。主成分分析表明，耳石总长 (TSL) 和最大宽度 (MW) 可作为表征春、夏产卵群体耳石长度和宽度的指标。春、夏产卵群体TSL生长模型采样年间差异均显著 (P<0.05)，春季产卵群体分别以逻辑斯蒂和对数生长模型拟合最佳，夏季产卵群体以逻辑斯蒂和线性拟合最佳。春、夏产卵群体MW生长模型采样年间差异均不显著 (P>0.05)，以逻辑斯蒂生长模型拟合最佳，性别间差异不显著 (P>0.05)。180~240 d，春季产卵群TSL和MW的生长速率均大于夏季产卵群体。Abstract: Based on Uroteuthis edulis samples collected from September 2017 to March 2018 and from September 2018 to March 2019, we observed the statolith morphology, and studied the microstructure and growth characteristics of the otolith. The results show that the statolith growth increments was composed of light and dark rings, and could be divided as postnuclear zone, dark zone and peripheral zone according to their width and brightness. According to the statolith age, the spawning population in spring and summer was the dominant cohort, and the morphological variables of the statolith between the two cohorts were significantly different (P<0.05). Principal component analysis shows that the total length (TSL) and maximum width (MW) of the otoliths could be used as indicators to characterize the length and width of statoliths in spring and summer spawning cohorts. The TSL growth function of the spring and summer spawning cohorts was significantly different in the two years (P<0.05). The former best fitted with Logistic and logarithmic growth functions, and the latter best fitted with Logistic and linear functions. The MW growth function of the spawning population in spring and summer showed no significant difference (P>0.05). The logistic growth function was the best fit and the gender difference was not significant (P>0.05). During the 180~240 days, the growth rate of spring and summer spawning cohorts both showed a downward trend, and the growth rates of TSL and MW in spring spawning cohorts were greater than those in summer.
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Key words:
- Uroteuthis edulis /
- statolith /
- Microstructure /
- Growth characteristics /
- East China Sea
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图 3 耳石形态分区及研磨平面
Figure 3. Morphometric dome of statolith and grinding plane of U. edulis
图 5 剑尖枪乌贼日龄及孵化日期分布
Figure 5. Distribution of age and hatching month for U. edulis in East China Sea
图 6 春季产卵群体耳石形态参数生长速率
Figure 6. Growth rate of morphological variables of otolith of spring hatching cohort
图 7 夏季产卵群体耳石形态参数生长速率
Figure 7. Growth rate of otolith morphological variables of summer hatching cohort
表 1 剑尖枪乌贼采样信息
Table 1. Sampling information of U. edulis in East China Sea
采样时间
Sampling time样本量
Number/尾胴长分布
Mantle range/mm2017年9月 September 2017 65 69~198 2017年10月 October 2017 28 75~135 2017年11月 November 2017 40 110~234 2017年12月 December 2017 49 86~223 2018年1月 January 2018 61 93~180 2018年2月 February 2018 30 70~158 2018年3月 March 2018 40 92~154 2018年9月 September 2018 101 62~198 2018年10月 October 2018 78 74~223 2018年11月 November 2018 30 95~170 2018年12月 December 2018 36 86~222 2019年1月 January 2019 44 60~227 2019年2月 February 2019 36 64~143 2019年3月 March 2019 80 58~164 
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表 2 剑尖枪乌贼耳石形态参数范围
Table 2. Range of statolith morphological variables of U. edulis
种群
Population形态变量
Statolith morphological variable2017 2018 最大值
Maximum最小值
Minimum平均值
Average最大值
Maximum最小值
Minimum平均值
Average春季产卵群体
Spring cohort of hatching耳石总长 TSL 1 844.85 1 235.61 1 668.59 1 801.94 1 351.45 1 596.00 翼区长 WL 1 497.33 1 059.71 1 278.37 1 402.94 986.78 1 221.98 吻区长 RL 823.75 420.47 659.79 819.45 467.82 636.64 最大宽度 MW 1 413.10 785.13 1 180.76 1 394.36 849.50 1 110.71 吻区宽 RW 499.52 326.07 419.12 471.94 321.89 392.38 翼区宽 WW 671.52 300.35 510.79 665.01 248.84 475.27 背侧区长 DLL 1 159.45 763.83 1 009.71 1 161.53 731.33 940.48 吻侧区长 RLL 1 550.46 1 007.74 1 336.71 1 522.09 1 121.98 1 313.07 侧区长 LDL 1 266.43 827.60 1 069.12 1 159.16 883.07 1 014.24 夏季产卵群体
Summer cohort of hatching耳石总长 TSL 1 737.59 1 394.36 1 572.81 1 754.75 1 372.90 1 513.06 翼区长 WL 1 308.56 1 003.94 1 166.24 1 385.78 995.36 1 163.41 吻区长 RL 733.66 467.65 621.41 798.05 484.83 583.37 最大宽度 MW 1 240.19 905.27 1 030.70 1 209.87 862.37 993.33 吻区宽 RW 467.67 227.55 351.13 420.45 291.74 342.76 翼区宽 WW 565.67 373.28 469.41 579.19 317.51 425.59 背侧区长 DLL 1 099.44 763.87 885.55 1 010.42 721.09 853.36 吻侧区长 RLL 1 391.98 1 090.39 1 256.22 1 468.65 1 080.02 1 254.73 侧区长 LDL 1 095.17 906.14 985.01 1 065.45 868.01 960.55
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表 3 剑尖枪乌贼耳石形态参数的4个主成分负荷值和贡献率
Table 3. Loadings of four principal components for statolith morphological of variables U. edulis
年份
Year形态变量
Statolith morphological variable春季产卵群体-主成分
Principal component ofspring hatching夏季产卵群体-主成分
Principal component of summer hatching1 2 3 4 1 2 3 4 2017 耳石总长 TSL 0.97 0.02 0.01 −0.01 0.89 −0.12 −0.22 −0.17 翼区长 WL 0.82 0.31 −0.23 −0.36 0.84 −0.40 −0.07 0.16 吻区长 RL 0.75 −0.58 0.06 −0.20 0.77 −0.20 −0.30 0.16 最大宽度 MW 0.96 −0.05 0.03 0.07 0.83 0.18 0.40 0.02 吻区宽 RM 0.67 −0.26 −0.52 0.43 0.47 0.62 −0.03 0.45 翼区宽 WM 0.66 −0.19 0.66 0.13 0.54 −0.48 0.52 −0.12 背侧区长 DLL 0.93 −0.15 0.00 0.06 0.76 0.20 0.45 0.13 吻侧区长 RLL 0.91 0.25 −0.09 −0.21 0.79 −0.04 −0.52 0.00 侧区长 LDL 0.58 0.73 0.17 0.27 0.60 0.47 −0.05 −0.63 贡献率 Contribution rate/% 66.75 12.85 8.90 5.45 54.09 12.53 11.60 7.87 累计贡献率 Cumulative contribution rate/% 66.75 79.60 88.50 93.95 54.09 66.62 78.22 86.10 2018 耳石总长 TSL 0.97 −0.08 −0.02 −0.07 0.96 −0.09 −0.10 0.16 翼区长 WL 0.85 −0.37 0.16 −0.18 0.89 −0.23 −0.28 0.11 吻区长 RL 0.80 −0.46 −0.09 0.24 0.88 −0.17 0.27 0.24 最大宽度 MW 0.94 0.18 −0.09 0.09 0.92 0.27 0.14 −0.11 吻区宽 RM 0.67 0.30 0.53 0.38 0.72 −0.41 0.43 −0.26 翼区宽 WM 0.71 0.24 −0.61 0.08 0.48 0.81 0.14 0.21 背侧区长 DLL 0.90 0.18 −0.09 0.15 0.88 0.31 0.12 −0.19 吻侧区长 RLL 0.90 −0.27 0.10 −0.18 0.87 −0.40 −0.17 0.11 侧区长 LDL 0.75 0.41 0.15 −0.46 0.78 0.23 −0.46 −0.27 贡献率 Contribution rate/% 70.38 9.04 8.22 5.77 69.27 14.29 7.09 3.84 累计贡献率 Cumulative contribution rate/% 70.38 79.42 87.64 93.41 69.27 83.56 90.65 94.49
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表 4 春、夏季产卵群体耳石总长和最大宽度生长模型的参数及赤池信息量比较
Table 4. Comparison of parameters and AIC for TSL and MW growth models of spring and summer hatching cohort
种群
Population形态变量
Morphological variable年份
Year生长模型
Growth pattern模型参数 Model parameters a b c R2 AIC 春季产卵群体
Spring cohort of hatching耳石总长 TSL 2017 线性 line 1 150.94 2.39 0.30 369.33 指数 Exponential 1 232.41 0.003 0.29 369.69 对数 Logarithm 536.71 −1216.58 0.31 368.10 幂函数 Power 307.49 0.32 0.31 368.50 逻辑斯蒂 Logistic 1 718.77 0.06 147.68 0.43 360.60 von Bertalanffy 1 673.35 −0.002 9 536.20 0.21 383.95 2018 线性 line 957.28 3.10 0.53 726.01 指数 Exponential 1 075.81 0.002 0.52 726.77 对数 Logarithm 649.40 −1861.60 0.53 724.64 幂函数 Power 186.10 0.40 0.53 725.10 逻辑斯蒂 Logistic 3 495.23 0.003 256.28 0.52 726.18 von Bertalanffy 1 595.32 −0.002 9 718.77 0.47 791.55 最大宽度 MW 2017—2018 线性 line 419.87 3.40 0.47 1137.5 指数 Exponential 617.92 0.003 0.45 1140.7 对数 Logarithm 731.21 −2773.46 0.49 1132.6 幂函数 Power 39.57 0.628 0.47 1135.6 逻辑斯蒂 Logistic 4 731.12 0.004 510.74 0.46 1139.7 Von Bertalanffy 1 133.30 −0.002 7 363.71 0.38 1219.0 夏季产卵群体
Summer cohort of hatching耳石总长 TSL 2017 线性 line 1 029.26 2.75 0.27 461.47 指数 Exponential 1 118.80 0.002 0.27 461.77 对数 Logarithm 553.00 −1349.29 0.28 460.63 幂函数 Power 251.26 0.35 0.30 460.92 逻辑斯蒂 Logistic 1 607.02 0.09 149.93 0.35 454.81 von Bertalanffy 1 571.76 −0.002 9 280.15 0.18 479.48 2018 线性 line 818.10 3.73 0.57 289.79 指数 Exponential 960.12 0.002 0.57 289.85 对数 Logarithm 700.44 −2146.00 0.57 289.92 幂函数 Power 135.23 0.46 0.57 289.81 逻辑斯蒂 Logistic 3 519.15 0.004 251.78 0.57 289.80 von Bertalanffy 1 521.39 −0.002 9 280.15 0.48 319.92 最大宽度 MW 2017—2018 线性 line 520.24 2.54 0.42 705.25 指数 Exponential 624.50 0.002 0.42 705.75 对数 Logarithm 489.73 −1 564.90 0.43 704.55 幂函数 Power 79,32 0.48 0.42 704.83 逻辑斯蒂 Logistic 4 108.58 0.003 531.77 0.42 705.58 von Bertalanffy 1 013.12 −0.002 7 511.95 0.39 756.07 注：下划线代表选择的最适宜生长模型
Note: The underlines represent fitted growth model.
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