植物生态学报 ›› 2022, Vol. 46 ›› Issue (9): 1050-1063.DOI: 10.17521/cjpe.2021.0463
哈里布努尔1, 古丽扎尔·阿不都克力木1,2, 热依拉穆·麦麦提吐尔逊1, 艾沙江·阿不都沙拉木1,2,*()
收稿日期:
2021-12-09
接受日期:
2022-05-07
出版日期:
2022-09-20
发布日期:
2022-10-19
通讯作者:
艾沙江·阿不都沙拉木
作者简介:
(aysajanxj@sina.com)基金资助:
Halibunuer 1, Gulzar ABDUKIRIM1,2, Reyilamu MAIMAITITUERXUN1, Aysajan ABDUSALAM1,2,*()
Received:
2021-12-09
Accepted:
2022-05-07
Online:
2022-09-20
Published:
2022-10-19
Contact:
Aysajan ABDUSALAM
Supported by:
摘要:
黑果枸杞(Lycium ruthenicum)是中国西北地区极端环境中分布的国家二级保护植物, 该物种在新疆南部的自然种群中出现了同型花柱类型(同位花)和柱头探出式雌雄异位类型(异位花)个体, 并且遭遇沙尘暴频繁的种群中异位花个体出现频率减少。该研究对喀什市自然种群中黑果枸杞两种不同花型植株的花部综合征和传粉特性进行比较研究, 以期探讨该物种不同花型植株在南疆早春极端环境中的花部特征的可塑性及其适应性机制。结果表明: 同位花雌雄蕊高度间无显著差异, 而异位花雌蕊高度显著高于雄蕊; 同位花花冠直径、花冠筒长、胚珠数均高于异位花, 而异位花雌雄蕊空间距离、花粉数及花粉胚珠比均比同位花高。黑果枸杞同位花个体比例(68%)高于异位花个体(32%), 种群水平及个体水平同位花花期((117.00 ± 2.25) d, (101.65 ± 1.98) d)比异位花((26.00 ± 1.00) d, (18.75 ± 1.00) d)长, 而单花水平上异位花单花寿命((4.50 ± 0.14) d)比同位花((3.13 ± 0.11) d)长。两种类型花在花早期(紫色)分泌的花蜜量均高于花后期(白色)。在紫色花阶段(花开放早期), 同位花上的主要传粉者意大利蜜蜂、熊蜂和食蚜蝇的访花频率和停留时间均高于异位花; 而白色花阶段(花开放后期)意大利蜜蜂、熊蜂在异位花上的访花频率比同位花高。在不同花色阶段, 同位花柱头花粉落置数、花粉移出率、花粉传递效率均比异位花高, 并且同位花自然坐果率及结籽率均比异位花高。在新疆南部的沙尘暴极端环境下, 同位花通过较高的自交亲和性保障繁殖, 而异交为主的异位花提高了异交率。异位花与同位花在花部综合征和花报酬上的差异, 是影响其繁殖成功的主要因素。
哈里布努尔, 古丽扎尔·阿不都克力木, 热依拉穆·麦麦提吐尔逊, 艾沙江·阿不都沙拉木. 黑果枸杞两种花型的花部综合征与传粉特性. 植物生态学报, 2022, 46(9): 1050-1063. DOI: 10.17521/cjpe.2021.0463
Halibunuer , Gulzar ABDUKIRIM, Reyilamu MAIMAITITUERXUN, Aysajan ABDUSALAM. Flower syndrome and pollination characteristics of two flower morphs in Lycium ruthenicum (Solanaceae). Chinese Journal of Plant Ecology, 2022, 46(9): 1050-1063. DOI: 10.17521/cjpe.2021.0463
图1 黑果枸杞花各部位特征测量(A)和不同花型花花柱及花药相对高度(B)(平均值±标准误)。a(a′)-e(e′), 花冠筒长度; a(a′)-d(d′), 雌蕊长; a(a′)-c(c′), 雄蕊长; f(f′)-b(b′), 花冠口直径。B中不同小写字母表示同一花型花不同性器官间差异显著(p < 0.05)。
Fig. 1 Measurements of flower traits (A) and relative height of sexual organs (B) of two flower morphs of Lycium ruthenicum (mean ± SE). a(a′)-e(e′), flower length; a(a′)-d(d′), pistil length; a(a′)-c(c′), stamen length; f(f′)-b(b′), corolla diameter. Different lowercase letters in B indicate significant difference between different sexual organs of the same flower morph (p < 0.05).
图2 新疆喀什黑果枸杞的植株、花序、果实、花型及花颜色变化。A, 雌雄异位花。B, 同位花。C, 异位花花序。D, 同位花花序。E, F, 果实。G, 植株。H, 不同花型花在花寿命内的花色变化(上部为同位花, 下部为异位花)。
Fig. 2 Individual plant, inflorescence, fruit, flower morphs and flower color changes in Lycium ruthenicum in Kashi, Xinjiang, China. A, Herkogamous flower. B, Homostylous flower. C, Inflorescence of herkogamous flower. D, Inflorescence of homostylous flower. E, F, Fruit. G, Individual plant. H, Change of flower color in different phases of flower longevity in two flower morphs of L. ruthenicum (upper, homostylous flower; lower, herkogamous flower).
花部特征 Flower characteristic | 紫色阶段 Purple phase | 白色阶段 White phase | 花型×阶段 Morph × phase | |||||||
---|---|---|---|---|---|---|---|---|---|---|
同位花 Homostylous flower | 雌雄异位花 Herkogamous flower | Wald χ12 | p1 | 同位花 Homostylous flower | 雌雄异位花 Herkogamous flower | Wald χ22 | p2 | Wald χ32 | p3 | |
花冠直径 Corolla diameter (mm) | 8.29 ± 0.25 | 7.36 ± 0.33 | 10.411 | 0.001 | 7.18 ± 0.23 | 6.70 ± 0.43 | 0.914 | 0.659 | 17.380 | 0.001 |
花冠筒长 Corolla tube length (mm) | 14.28 ± 0.31 | 10.70 ± 0.54 | 32.514 | <0.001 | 14.92 ± 0.39 | 9.71 ± 0.57 | 57.284 | <0.001 | 4.188 | 0.242 |
雌蕊长 Pistil length (mm) | 11.99 ± 0.33 | 12.20 ± 0.57 | 0.104 | 0.747 | 12.06 ± 0.33 | 13.07 ± 0.48 | 3.005 | 0.083 | 0.043 | 0.998 |
雄蕊长 Stamen length (mm) | 11.99 ± 0.24 | 10.20 ± 0.42 | 13.574 | <0.001 | 12.37 ± 0.36 | 11.14 ± 0.53 | 3.745 | 0.053 | 1.481 | 0.687 |
雌雄蕊空间距离 Stamen-stigma distance (mm) | 0.65 ± 0.16 | 2.00 ± 0.27 | 18.588 | <0.001 | 1.02 ± 0.18 | 1.93 ± 0.27 | 8.042 | 0.005 | 4.358 | 0.225 |
雌蕊-花冠口距离 Corolla tube-stigma distance (mm) | 2.29 ± 0.22 | 1.50 ± 0.39 | 3.175 | 0.075 | 2.86 ± 0.28 | 3.36 ± 0.42 | 0.975 | 0.323 | 12.430 | 0.006 |
雄蕊-花冠口距离 Corolla tube-stamen distance (mm) | 0.70 ± 0.28 | 2.29 ± 0.16 | 24.444 | <0.001 | 2.55 ± 0.29 | 1.86 ± 0.42 | 1.865 | 0.172 | 1.208 | 0.751 |
表1 黑果枸杞同位花和雌雄异位花在花寿命早期(紫色)和后期(白色)花部特征(平均值±标准误)的比较(广义线性模型)
Table 1 Comparisons of flower traits (mean ± SE) among two flower morphs at early flowering phase (purple) and late flowering phase (white) of Lycium ruthenicum using generalized linear model (GLM)
花部特征 Flower characteristic | 紫色阶段 Purple phase | 白色阶段 White phase | 花型×阶段 Morph × phase | |||||||
---|---|---|---|---|---|---|---|---|---|---|
同位花 Homostylous flower | 雌雄异位花 Herkogamous flower | Wald χ12 | p1 | 同位花 Homostylous flower | 雌雄异位花 Herkogamous flower | Wald χ22 | p2 | Wald χ32 | p3 | |
花冠直径 Corolla diameter (mm) | 8.29 ± 0.25 | 7.36 ± 0.33 | 10.411 | 0.001 | 7.18 ± 0.23 | 6.70 ± 0.43 | 0.914 | 0.659 | 17.380 | 0.001 |
花冠筒长 Corolla tube length (mm) | 14.28 ± 0.31 | 10.70 ± 0.54 | 32.514 | <0.001 | 14.92 ± 0.39 | 9.71 ± 0.57 | 57.284 | <0.001 | 4.188 | 0.242 |
雌蕊长 Pistil length (mm) | 11.99 ± 0.33 | 12.20 ± 0.57 | 0.104 | 0.747 | 12.06 ± 0.33 | 13.07 ± 0.48 | 3.005 | 0.083 | 0.043 | 0.998 |
雄蕊长 Stamen length (mm) | 11.99 ± 0.24 | 10.20 ± 0.42 | 13.574 | <0.001 | 12.37 ± 0.36 | 11.14 ± 0.53 | 3.745 | 0.053 | 1.481 | 0.687 |
雌雄蕊空间距离 Stamen-stigma distance (mm) | 0.65 ± 0.16 | 2.00 ± 0.27 | 18.588 | <0.001 | 1.02 ± 0.18 | 1.93 ± 0.27 | 8.042 | 0.005 | 4.358 | 0.225 |
雌蕊-花冠口距离 Corolla tube-stigma distance (mm) | 2.29 ± 0.22 | 1.50 ± 0.39 | 3.175 | 0.075 | 2.86 ± 0.28 | 3.36 ± 0.42 | 0.975 | 0.323 | 12.430 | 0.006 |
雄蕊-花冠口距离 Corolla tube-stamen distance (mm) | 0.70 ± 0.28 | 2.29 ± 0.16 | 24.444 | <0.001 | 2.55 ± 0.29 | 1.86 ± 0.42 | 1.865 | 0.172 | 1.208 | 0.751 |
图3 黑果枸杞不同花型花紫色阶段(1、2和3)和白色阶段(4和5)花粉活力及柱头可授性比较(平均值±标准误)。“+”表示柱头可授性强弱程度。
Fig. 3 Pollen viability and stigma acceptability in purple phase (1, 2 and 3) and white phase (4 and 5) of two flower morphs of Lycium ruthenicum (mean ± SE). “+” indicates the degree of acceptability of stigma.
图4 黑果枸杞不同花型花在花寿命早期(紫色)和后期(白色)的花蜜分泌量(平均值±标准误)。不同大写字母表示同种类型不同阶段间的差异显著, 不同小写字母表示同一阶段不同花型间的差异显著(p < 0.05)。
Fig. 4 Nectar volume at early flowering phase (purple) and late phase (white) of two flower morphs of Lycium ruthenicum (mean ± SE). Different uppercase letters indicate significant difference between different phases of the same flower morph, and different lowercase letters indicate significant difference between different flower morphs at the same phase (p < 0.05).
传感器 Sensor | 物质 Substance | 紫色阶段 Purple phase | 白色阶段 White phase | 花型×阶段 Morph × phase | |||||||
---|---|---|---|---|---|---|---|---|---|---|---|
同位花 Homostylous flower | 雌雄异位花 Herkogamy flower | Wald χ12 | p1 | 同位花 Homostylous flower | 雌雄异位花 Herkogamy flower | Wald χ22 | p2 | Wald χ32 | p3 | ||
W1C | 芳香成分苯类 Benzenoids | 8.58 ± 0.37 | 9.26 ± 0.46 | 1.408 | 0.235 | 6.31 ± 0.20 | 8.23 ± 0.04 | 130.437 | <0.001 | 3.385 | 0.066 |
W3C | 氨类, 芳香成分 Ammonia, benzenoids | 8.19 ± 0.40 | 7.01 ± 1.49 | 0.463 | 0.496 | 5.45 ± 0.27 | 7.76 ± 0.05 | 106.682 | <0.001 | 3.120 | 0.077 |
W5C | 烷烃芳香成分 Alkane aromatic composition | 8.06 ± 0.41 | 8.92 ± 0.55 | 1.566 | 0.211 | 5.28 ± 0.28 | 7.64 ± 0.05 | 103.141 | <0.001 | 3.424 | 0.064 |
W2W | 芳香成分, 有机硫化物 Benzenoids, organic sulfide | 10.92 ± 0.39 | 10.11 ± 0.27 | 4.025 | 0.045 | 14.97 ± 0.24 | 10.28 ± 0.22 | 309.736 | <0.001 | 87.539 | <0.001 |
W1S | 甲基类 Methyl | 19.83 ± 0.55 | 15.87 ± 1.60 | 4.450 | 0.035 | 25.47 ± 1.01 | 19.86 ± 0.31 | 42.106 | <0.001 | 0.531 | 0.466 |
W1W | 无机硫化物 Inorganic sulfide | 7.61 ± 2.61 | 10.25 ± 0.22 | 2.484 | 0.115 | 13.67 ± 0.19 | 10.48 ± 0.14 | 267.990 | <0.001 | 9.389 | 0.002 |
W2S | 醇类, 醛酮类 Alcohols, aldehydes and ketones | 10.40 ± 0.30 | 10.67 ± 0.14 | 1.223 | 0.269 | 10.92 ± 0.39 | 10.70 ± 0.10 | 0.357 | 0.550 | 0.279 | 0.598 |
W3S | 烷烃 Alkane | 8.28 ± 0.40 | 8.99 ± 0.53 | 1.131 | 0.288 | 5.37 ± 0.27 | 7.77 ± 0.04 | 112.877 | <0.001 | 4.695 | 0.030 |
W5S | 氮氧化合物 Nitrogen oxides | 9.68 ± 0.29 | 9.85 ± 0.33 | 0.160 | 0.689 | 7.41 ± 0.36 | 9.35 ± 0.09 | 40.361 | <0.001 | 10.389 | 0.001 |
W6S | 氢化物 Hydride | 8.46 ± 0.39 | 9.08 ± 0.51 | 0.952 | 0.329 | 5.46 ± 0.26 | 7.93 ± 0.05 | 125.947 | <0.001 | 6.120 | 0.013 |
表2 新疆喀什黑果枸杞不同花型花在花寿命不同阶段(紫色和白色)花气味中的化学成分占总化合物百分比(平均值±标准误)的比较
Table 2 Mean relative composition of two flower morphs at early flowering phase (purple) and late flowering phase (white) of Lycium ruthenicum flower volatiles in Kashi, Xinjiang, China (mean ± SE)
传感器 Sensor | 物质 Substance | 紫色阶段 Purple phase | 白色阶段 White phase | 花型×阶段 Morph × phase | |||||||
---|---|---|---|---|---|---|---|---|---|---|---|
同位花 Homostylous flower | 雌雄异位花 Herkogamy flower | Wald χ12 | p1 | 同位花 Homostylous flower | 雌雄异位花 Herkogamy flower | Wald χ22 | p2 | Wald χ32 | p3 | ||
W1C | 芳香成分苯类 Benzenoids | 8.58 ± 0.37 | 9.26 ± 0.46 | 1.408 | 0.235 | 6.31 ± 0.20 | 8.23 ± 0.04 | 130.437 | <0.001 | 3.385 | 0.066 |
W3C | 氨类, 芳香成分 Ammonia, benzenoids | 8.19 ± 0.40 | 7.01 ± 1.49 | 0.463 | 0.496 | 5.45 ± 0.27 | 7.76 ± 0.05 | 106.682 | <0.001 | 3.120 | 0.077 |
W5C | 烷烃芳香成分 Alkane aromatic composition | 8.06 ± 0.41 | 8.92 ± 0.55 | 1.566 | 0.211 | 5.28 ± 0.28 | 7.64 ± 0.05 | 103.141 | <0.001 | 3.424 | 0.064 |
W2W | 芳香成分, 有机硫化物 Benzenoids, organic sulfide | 10.92 ± 0.39 | 10.11 ± 0.27 | 4.025 | 0.045 | 14.97 ± 0.24 | 10.28 ± 0.22 | 309.736 | <0.001 | 87.539 | <0.001 |
W1S | 甲基类 Methyl | 19.83 ± 0.55 | 15.87 ± 1.60 | 4.450 | 0.035 | 25.47 ± 1.01 | 19.86 ± 0.31 | 42.106 | <0.001 | 0.531 | 0.466 |
W1W | 无机硫化物 Inorganic sulfide | 7.61 ± 2.61 | 10.25 ± 0.22 | 2.484 | 0.115 | 13.67 ± 0.19 | 10.48 ± 0.14 | 267.990 | <0.001 | 9.389 | 0.002 |
W2S | 醇类, 醛酮类 Alcohols, aldehydes and ketones | 10.40 ± 0.30 | 10.67 ± 0.14 | 1.223 | 0.269 | 10.92 ± 0.39 | 10.70 ± 0.10 | 0.357 | 0.550 | 0.279 | 0.598 |
W3S | 烷烃 Alkane | 8.28 ± 0.40 | 8.99 ± 0.53 | 1.131 | 0.288 | 5.37 ± 0.27 | 7.77 ± 0.04 | 112.877 | <0.001 | 4.695 | 0.030 |
W5S | 氮氧化合物 Nitrogen oxides | 9.68 ± 0.29 | 9.85 ± 0.33 | 0.160 | 0.689 | 7.41 ± 0.36 | 9.35 ± 0.09 | 40.361 | <0.001 | 10.389 | 0.001 |
W6S | 氢化物 Hydride | 8.46 ± 0.39 | 9.08 ± 0.51 | 0.952 | 0.329 | 5.46 ± 0.26 | 7.93 ± 0.05 | 125.947 | <0.001 | 6.120 | 0.013 |
图5 黑果枸杞主要访花昆虫。A, 意大利蜜蜂。B, 熊蜂。C, 食蚜蝇。D, 淡脉隧蜂。E, 丝光绿蝇。
Fig. 5 Main visiting insects of Lycium ruthenicum. A, Apis mellifera. B, Bombus sp. C, Syrphidae. D, Lasioglossum occidens. E, Lucilia sericata.
图6 黑果枸杞不同花型花主要传粉者在花寿命早期(紫色)和后期(白色)的访花频率及停留时间(平均值±标准误)。不同大写字母表示同种花型不同传粉者间的差异, 不同小写字母表示同一传粉者不同花型间的差异显著(p < 0.05)。
Fig. 6 Visiting frequency of main pollinators at early flowering phase (purple) and late flowering phase (white) of Lycium ruthenicum of two flower morphs (mean ± SE). Different uppercase letters indicate significant difference between different pollinators of the same morph, and different lowercase letters indicate significant difference between different flower morphs of same pollinator (p < 0.05).
图7 黑果枸杞不同花型花在花寿命早期(紫色)和晚期(白色)花粉移出率(A)、柱头落置花粉数(B)、花粉传递效率(C)、自然坐果率和结籽率(D)(平均值±标准误)。A-C不同大写字母表示同种花型不同阶段间的差异显著, 不同小写字母表示不同花型间差异显著(p < 0.05)。
Fig. 7 Proportion of total pollen grains removed (A), deposited pollen grains (B), pollen transfer efficiency (C) and fruit and seed set rate (D) in the two flower morphs of Lycium ruthenicum at early flowering phase (purple) and late flowering phase (white)(mean ± SE). Different uppercase letters indicate significant difference between different phases of the same morph, and different lowercase letters indicate significant difference between different flower morph at the same phase (p < 0.05).
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