不同世代生姜组培苗物质生产特性与产量形成差异

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任云, 李强, 李哲馨, 刘奕清, 唐建民. 不同世代生姜组培苗物质生产特性与产量形成差异[J]. 浙江农业学报, 2020,32(4): 661-670.
REN Yun, LI Qiang, LI Zhexin, LIU Yiqing, TANG Jianmin. Differences in matter production characteristics and yield formation of tissue culture gingers in different generations[J]. ACTA AGRICULTURAE ZHEJIANGENSIS, 2020,32(4): 661-670.
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不同世代生姜组培苗物质生产特性与产量形成差异

任云, 李强, 李哲馨, 刘奕清, 唐建民^*

重庆经济植物生物技术重点实验室,重庆市特种植物协调创新中心,重庆文理学院特色植物研究院,重庆 402160

*通信作者,唐建民,E-mail: 756337153@qq.com

作者简介:任云(1989—),女,湖南常德人,博士,讲师,主要从事园艺植物逆境生理调控。E-mail: 839953291@qq.com

收稿日期: 2010-10-25

基金资助: 重庆市教委自然科学基金青年基金(KJQN201801330,KJQN201801339); 重庆文理学院科研项目(2017RTZ18)

摘要

为研究不同世代组培生姜物质积累与分配特性及其产量形成差异,以确定组培生姜在生产上的最佳推广世代。以组培生姜原原种(T₁)、原种(T₂)、生产种(T₃)3个世代为试验材料,于2017—2018年进行了大田栽培试验。结果表明:T₃和T₂世代植株生长特性优于T₁,表现为生育前期T₃和T₂株高、茎粗快速增加,地上部快速生长;进入根茎膨大期后,生姜株高和茎粗增长减缓,但分蘖快速发生,由于T₃和T₂的分蘖发生少,保证了根茎光合产物的供给,延长了快速生长期,显著提高了其物质积累,为高产奠定了物质基础。根茎膨大期后,T₃和T₂较低的茎、叶和根系物质分配比例,使光合产物向根茎转移促进了根茎的膨大,从而获得高产。因此,与原原种和原种相比,生姜组培生产种具有明显的产量优势,能够有效提高生姜生产的经济效益。综合考虑不同世代组培生姜产量和经济效益差异与种性退化规律,可以把组培生姜生产种(T₃)作为生产上的最佳推广世代,既能充分发挥其产量潜力提高经济效益,又能有效控制病害的发生。

关键词: 生姜; 世代; 组培; 物质积累与分配; 产量

中图分类号:S632.5 文献标志码:A 文章编号:1004-1524(2020)04-0661-10

Differences in matter production characteristics and yield formation of tissue culture gingers in different generations

REN Yun, LI Qiang, LI Zhexin, LIU Yiqing, TANG Jianmin^*

Chongqing Key Laboratory of Economic Plant Biotechnology, Collaborative Innovation Center of Special Plant Industry in Chongqing, Institute of Special Plants, Chongqing University of Arts and Sciences, Chongqing 402160, China

Abstract

To study the differences in matter production characteristics and yield formation of tissue cultured gingers in different generations, and to determine the best promotion generation of tissue cultured ginger in production, tissue cultured original-original seeds (T₁), original seeds (T₂) and production seeds (T₃) were used as experimental materials to conduct a two-year field experiment in 2017-2018. The results showed that the growth characteristics of T₃ and T₂ were significantly better than that of T₁, which showed the plant height and stem diameter of T₃ and T₂ grew rapidly, and ensured the rapid growth of shoot in the early growth stage. After entering the rhizome expansion period, the growth of plant height and stem diameter of ginger slowed down, and tillers occurred rapidly. T₃ and T₂ could effectively control the occurrence of tillering, reduce the consumption of photosynthetic products by excessive tillering, and ensure the supply of photosynthetic products of rhizomes, thereby increased the rate of dry matter accumulation, prolonged the rapid growth period, and significantly increased their dry matter accumulation, laid a material foundation for high yield. After rhizome expansion period, the lower proportion in stem, leaf and root of T₃ and T₂, which promoted the transfer of photosynthetic products to rhizomes and further promoted the expansion of rhizomes, thus achieved high yield. Therefore, compared with the original-original seeds and original seeds, the production seeds of ginger had obvious yield advantages, which could effectively improve the economic benefits of ginger production. Comprehensive considering the yield differences and economic benefits, and seeds degeneration law of tissue cultured ginger, the tissue cultured ginger production seeds (T₃) could be used as the best promotion generation for production, which could realize its yield potential, improve economic benefits and effectively control the occurrence of diseases.

Keyword: ginger; generation; tissue culture; matter accumulation and distribution; yield

文章图片

生姜(Zingier officinal Roscoe)为姜科多年生草本植物, 具有食用和药用双重价值, 在世界范围内广泛种植^{[1, 2]}。我国生姜在全球市场占主导地位, 栽培面积与总产量均居世界首位^[3]。同时, 生姜作为我国优势特色蔬菜和主要调味品原料, 已成为乡村振兴与脱贫攻坚的特色高效产业, 也是重庆市现代特色效益农业调味品产业体系主导作物之一^{[4, 5]}。

生姜以地下根茎无性繁殖为主, 连年自留种, 种性退化, 病原物积累, 土传病害严重, 使得生姜生长缓慢, 品质下降, 减产甚至绝收, 严重制约生姜产业的持续健康发展^{[6, 7]}。而组织培养是解决无性繁殖作物种性退化、病原物积累, 提高抗病性和产量潜力的重要措施^{[8, 9, 10, 11]}。王林生等^[12]研究表明, 不同世代组培甘薯增产幅度差异明显, 茎尖苗增产效果明显高于原原种苗、原种苗和良种苗。汤青林等^[13]通过测定不同世代组培芋苗叶片生理指标和产量, 综合评价指出组培2代要优于组培1代和普通苗, 是大面积推广种植的种源。组培技术在甘薯、马铃薯上已有大量研究应用^{[14, 15]}。生姜的组培多是关于其组培快繁体系建立的相关研究, 而关于不同世代组培生姜物质积累与分配特性, 以及产量形成差异的研究还鲜有报道。本研究以生姜原原种(T₁)、原种(T₂)、生产种(T₃)为试验材料, 研究了不同世代组培生姜物质生产特性与产量形成差异, 以期为生姜组培种生产上最佳推广世代的确定及其增产机制的研究提供理论依据。

1 材料与方法

1.1 试验地概况

试验于2017年和2018年在永川区五间镇试验基地进行, 该地区属于亚热带季风性湿润气候, 年平均温度17.7 ℃, 年平均降雨量1 015.0 mm, 日均日照时数3.3 h, 年均无霜期317 d。该地区为浅丘旱地, 以特色经济作物为主。2年试验均在同一地块进行, 2017年播种前试验地0~30 cm土层有机质含量18.7 g· kg^-1, 碱解氮41.6 mg· kg^-1, 速效磷122.1 mg· kg^-1, 速效钾338.8 mg· kg^-1, 全氮1.3 g· kg^-1, 全磷17.9 g· kg^-1, 全钾1.6 g· kg^-1, pH为6.84。

1.2 方法

试验生姜品种为竹根姜, 供试材料为本实验室自主培育的不同世代竹根姜脱毒姜种。原原种(T₁)为组培试管苗, 原种(T₂)为原原种种植收获的姜种, 生产种(T₃)为原种种植收获的姜种。

采用随机区组试验设计, 共设3个处理, 分别为原原种(T₁)、原种(T₂)、生产种(T₃), 3次重复, 小区面积50 m²。常规化肥施氮(N)、磷(P₂O₅)、钾(K₂O)用量分别为400、200、600 kg· hm^-2, 有机肥1 500 kg· hm^-2。磷肥、钾肥和有机肥一次性基施, 氮肥50%基施, 50%块茎膨大期追肥。种植规格为畦带沟宽1.8 m, 双行种植, 株行距为0.5 m× 0.3 m, 种植密度为90 000 株· hm^-2, 其他栽培管理同当地生姜高产田。2017年于4月28日播种, 11月20日收获; 2018年于5月1日播种, 11月25日收获计产。

1.3 指标测定

1.3.1 形态指标测定

生姜出苗后, 每隔20 d每小区选取连续10株长势一致的生姜植株, 测定株高、茎粗、分蘖数。株高测定从主茎基部量到生姜植株顶部; 茎粗采用电子游标卡尺测量每一株生姜茎粗最大的分蘖; 分蘖数采用直接计数法。

1.3.2 植物各器官干物质积累与分配

于生姜主要生育阶段(2017年出苗后105、126、150、189和207 d, 2018年出苗后81、113、144、174和193 d)选取长势一致的生姜植株5株, 按根、块茎、茎鞘和叶各器官分开, 测定各部分干质量, 并计算生姜的根冠比和物质分配比例。

利用Logistic方程对生姜物质生产进行曲线拟合, 可以得出最大理论生物量(final theoretical biomass yield, M_D)、最大生长速率(maximum growth rate, V_mD)、平均生长速率(average growth rate, V_aD)、最大生长速率出现时间点(time of instantaneous maximum slope, T_0D)、生长加速点(time of growth rate acceleration, T_1D), 生长减速点(time of growth rate deceleration, T_2D)和快速生长期(rapid growth rate period, T_2D-T_1D)^{[16, 17, 18]}。

y= $\frac{M_{D}}{1 + a EX P^{- bx}}$ ;

V_mD= $\frac{Mb}{4}$ ;

V_aD= $\frac{M}{Growthperiod}$ ;

T₀= $\frac{\sqrt[]{a}}{b}$ ;

T₁_D= $\frac{lna - 1.317}{b}$ ;

T₂_D= $\frac{lna + 1.317}{b}$ 。

式中:x, 生长天数; y, 干物质积累量; a、EXP和b代表方程系数; m, 实际干物质积累量; T₀, 最大积累速率时间点。

1.3.3 产量测定

收获时, 小区实收计产, 并按小区面积折算不同处理生姜实际产量。

1.4 数据分析

利用SPSS 20.0统计软件进行数据统计分析, 用LSD法对数据进行差异显著性检验, 用GraphPad Prism 5.0进行绘图, 利用OriginPro 2016进行生姜生长曲线拟合。

2 结果与分析

2.1 不同世代组培生姜植株生长特性的差异

图1表明, 不同世代生姜组培苗株高均随生育进程而逐渐增加, 但不同世代生姜组培苗株高差异显著。2017年和2018年各世代株高均表现为T₃> T₂> T₁, 其中各时期T₃较T₂株高的增幅均明显高于T₂较T₁株高的增幅, T₃较T₂各时期平均株高增幅分别为55.74% (2017年)和55.49%(2018年), T₂较T₁各时期平均株高增幅分别为44.42%(2017年)和36.77%(2018年)。T₃较T₂和T₂较T₁株高的增幅均随生育进程而表现出先增加后降低的变化趋势, 出苗80 d左右, T₂较T₁株高增幅最大, 增幅达61.24%; T₃较T₂最大株高增幅也出现在出苗80 d左右, 增幅达68.53%。表明生产种(T₃)较原种(T₂)和原原种(T₁)具有明显的株高优势。

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	图1 不同世代组培生姜株高差异Fig.1 Plant height differences of different generation tissue culture gingers

不同世代组培生姜茎粗均表现出前期增长较快, 后期增长放缓, 但不同世代生姜茎粗差异显著, 2017和2018年各时期均表现出T₃> T₂> T₁的变化趋势(图2)。2017、2018年收获时T₂较T₁茎粗分别高19.84%和19.66%, 而T3较T2分别高17.35%和16.27%, 表明随组培世代的增加, 组培生姜茎粗增幅下降。2017、2018年T₂较T₁与T₃较T₂茎粗差异均表现为随生育进程逐渐下降。2017年T₂较T₁茎粗增幅从49.85%下降到19.84%, 2018年从28.22%下降到19.66%; 2017年T₃较T₂茎粗增幅从37.11%下降到17.35%, 2018年从46.70%下降到16.27%。表明原种(T₂)和生产种(T₃)生育前期能保持较好的形态建成, 保证其快速生长期的物质积累, 尤其是生产种。

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	图2 不同世代组培生姜茎粗差异Fig.2 Stem diameter differences of different generation tissue culture gingers

图3结果表明, 不同世代组培生姜分蘖数均表现为生育前期增幅缓慢, 后期快速增加, 出苗80 d以前各世代分蘖数均小于10; 出苗80 d以后, T₁、T₂和T₃世代生姜2年平均分蘖数增幅分别达3.66· (10d)^-1、2.56· (10d)^-1和1.97· (10d)^-1。2017、2018年成熟期分蘖数均表现为T₁> T₂> T₃, 随世代的增加分蘖数显著下降, 2017和2018年收获时T₂较T₁分蘖数分别下降29.90%和27.52%, T₃较T₂分蘖数分别下降13.42%和15.44%。表明原种(T₂)和生产种(T₃)在生育后期能够有效控制分蘖的产生, 减少过多分蘖对光合产物的消耗, 从而提高其产量。

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	图3 不同世代组培生姜分蘖数差异Fig.3 Tillering number differences of different generation tissue culture gingers

2.2 不同世代组培生姜干物质积累与分配差异

生姜整个生育期内物质积累呈“ S” 型增长曲线, 本研究拟合的所有Logistic方程的决定系数R²均大于0.99(图4)。不同世代组培生姜理论物质积累量(M_D)、最大积累速率(V_mD)、平均积累速率(V_aD)、最大积累速率出现时间点(T_0D)和快速增长期(T_2D-T_1D)差异均显著(图4和表1)。随世代的增加, M_D、V_mD、V_aD均表现出逐渐增加的趋势, T_0D有所提前, 使得T_2D-T_1D显著延长。2017、2018年T₂较T₁平均快速增长期延长了11.07%(5.85 d), T₃较T₁延长了45.80%(24.22 d)。因此, 随组培世代的增加, 干物质积累速率逐渐提高, 快速增长期逐渐延长, 显著提高了原种(T₂)和生产种(T₃)的物质积累量。T₂较T₁干物质积累量分别提高了27.32%(2017年)和22.77%(2018年), T₃较T₂分别提高了28.20%(2017年)和31.50%(2018年)。

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	图4 不同世代组培生姜干物质积累拟合曲线Fig.4 Dry matter accumulation fitting curves of different generation tissue culture gingers

表1 不同世代组培生姜物质积累特性差异 Table 1 Differences in matter accumulation characteristics of different generation tissue culture gingers

2年试验中, 生姜生育中后期其叶片和茎的干物质分配比例均表现为随生育进程推进而降低, 采收时最低; 而块茎和根的干物质分配比例均表现为随生长期延长而增加, 采收时比例最高(图5)。2017、2018年收获时叶片、茎和根的干物质分配比例均表现出随世代增加而降低的趋势, 均为T₁> T₂> T₃; 而块茎的分配比例则随世代增加而增加, 均为T₃> T₂> T₁。收获时2年平均叶片分配比例T₁较T₂、T₃分别高4.50和10.73百分点, 茎分配比例分别高3.80和5.18百分点, 根分配比例分别高1.72和6.95百分点, 而块茎分配比例T₁较T₂、T₃则分别低10.00和22.86百分点。各世代组培生姜块茎分配比例差异明显高于其他器官, 表明不同世代组培生姜块茎物质分配比例差异是引起其产量差异的重要原因。

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	图5 不同世代组培生姜物质分配差异Fig.5 Matter distribution differences of different generation tissue culture gingers

生姜生育期内各世代生姜根冠比均表现出逐渐升高的变化趋势, 在成熟期达到最大(图6)。生育前期生姜需要进行地上部的形态建成, 较低的根冠比可以保证地上部的旺盛生长, 而到生育后期生姜以地下块茎为收获对象, 根冠比越高越利于高产。随世代的增加, 生姜根冠比逐渐升高, 2年均表现为T₃> T₂> T₁, 收获时T₂较T₁分别高出37.55%(2017年)和42.39% (2018年), T₃较T₁分别高出93.26%(2017年)和87.38%(2018年)。因此, 较高的根冠比是原种(T₂)和生产种(T₃)高产的重要原因。

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	图6 不同世代组培生姜根冠比变化Fig.6 Changes of root-to-shoot ratio of different generation tissue culture gingers

2.3 不同世代组培生姜产量差异

不同世代组培生姜产量差异显著, 2017年、2018年均表现出T₃> T₂> T₁(图7)。T₂较T₁增产56.18%(2017年)和47.48%(2018年), 而T₃产量较T₁高出120.92%(2017年)和109.87%(2018年), 表明生产种(T₃)较原原种(T₁)和原种(T₂)在产量上均有较大的提高, 更能发挥生姜品种的产量潜力。

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	图7 不同世代组培生姜产量差异Fig.7 Yield differences of different generation tissue culture gingers

2.4 不同世代组培生姜生产效益分析

表2结果表明, 不同世代组培生姜生产的投入差异较大, 经济效益也存在显著差异。由于生姜组培苗价格较高, 所以2年姜种投入均表现出随世代增加而降低, T₁> T₂> T₃, 而肥料、农药与其他投入在处理之间差异不大, 使得总投入也表现出T₁> T₂> T₃的趋势。总收入和效益2年均为生产种(T₃)显著高于T₁和T₂, 而T₁与T₂差异较小, 2年平均T₃总收入较T₁、T₂分别高出34.21%和41.88%, 效益分别高出66.25%和66.12%。由于T₁总投入较高, 使得产投比2年均表现出T₃> T₂> T₁。

表2 不同世代组培生姜生产的投入与产出 Table 2 Input and output of different generation tissue culture ginger production yuan· h

{m^{-}}^{2}

年份 Year	处理 Treatment	效益 Efficiency	总投入 Total inputs	姜种 Seeds	肥、药 Fertilizer and pesticide	其他 Other	总收入 Total income	产投比 Output to input ratio
2017	T₁	217063	123900	54000	9900	60000	340963	2.75
	T₂	204728	105900	36000	9900	60000	310628	2.93
	T₃	341156	98250	28350	9900	60000	439406	4.47
2018	T₁	195004	123900	54000	9900	60000	318904	2.57
	T₂	207645	105900	36000	9900	60000	313545	2.96
	T₃	343887	102300	32400	9900	60000	446187	4.36

The tissue culture seedling were 0.6 yuan· plant^-1 and original seeds 12 yuan· kg^-1 in both 2017 and 2018. The production seeds was 7 yuan· kg^-1 in 2017 and 8 yuan· kg^-1 in 2018.

2017和2018年组培苗均为0.6 元· 株^-1, 原种均为12 元· kg^-1, 而生产种2017年7元· kg^-1, 2018年8元· kg^-1。

表2 不同世代组培生姜生产的投入与产出 Table 2 Input and output of different generation tissue culture ginger production yuan· h

{m^{-}}^{2}

3 结论与讨论

T₃和T₂生长特性明显优于T₁, 生育前期株高、茎粗快速生长, 保证其地上部快速生长; 进入根茎膨大期后, 生姜株高和茎粗的增长减缓, 分蘖快速发生, 能够有效地控制分蘖的发生, 减少分蘖过多对光合产物的消耗, 保证了根茎光合产物的供给, 从而提高其物质积累速率, 延长快速生长期, 并显著提高其干物质积累, 为高产奠定了物质基础。进入根茎膨大期后, 较低的茎、叶和根系物质分配比例, 促进光合产物向根茎的转移, 进一步促进根茎的膨大, 从而获得高产。因此, 与原原种和原种相比, 生姜组培生产种具有明显的产量优势, 能够有效提高生姜生产的经济效益。综合考虑不同世代组培生姜产量和经济效益差异与种性退化规律, 可以把组培生姜生产种(T₃)作为生产上的最佳推广世代, 既能充分发挥其产量潜力、提高经济效益, 又能有效控制病害的发生。

作物产量与植株形态特征密切相关, 茎基部较粗、蔓较短、分枝数较少、薯块大小均匀的甘薯往往产量较高^[19]。高产型生姜的主要特征为茎秆粗壮、分蘖较少、叶片较厚、同化系统发达、根茎干物质含量较低^{[5, 20]}。本研究结果表明, 不同世代组培生姜株高、茎粗和分蘖数差异显著, 株高与茎粗均随世代的增加而显著增加, 而分蘖数随世代增加而显著降低。生产种生长状况明显优于原种和原原种, 生育前期株高、茎粗快速生长, 保证其地上部快速生长, 能够更好地进行光合作用, 为后期根茎的生长奠定良好的物质基础。进入块茎膨大期后, 株高和茎粗的增长减缓, 分蘖快速发生, 生产种能够有效地控制分蘖的发生, 减少分蘖过多对光合产物的消耗, 保证了根茎光合产物的供给。因此, 株高、茎粗增加, 分蘖减少是生产种较原种和原原种增产的重要原因。

干物质积累与分配是作物产量形成的基础, 脱毒甘薯干物质的积累, 在整个生育期内均明显高于对照, 其分配规律为:前期以地上部茎叶为主, 中后期地下部块根迅速增加, 从而提高产量^{[11, 12]}。本研究结果表明, 生产种(T₃)干物质积累速率提高, 快速增长期延长, 各时期的物质积累量增加为其高产奠定了物质基础。生姜生育前期主要进行地上部形态建成, 物质分配以茎叶为主, 根冠比较低; 而进入根茎膨大期后, 物质积累从地上向地下转移, 根茎物质分配比和根冠比逐渐提高, 茎叶分配比则随之下降, 与王林生等^[12]在甘薯上的研究结果一致。组培生姜茎、叶片和根物质分配比例均随组培世代的增加而降低, 而块茎分配比例则随组培世代的增加而升高。原种和原原种较低的物质积累速率和较短的快速增长期, 使得其物质积累量显著低于生产种, 从根本生影响其产量形成; 进入根茎膨大期后, 茎叶较高的物质分配比影响了原种和原原种光合产物从地上部往根茎的转运, 进一步限制了根茎的膨大; 原种和原原种需要较多的根系才能满足其植株的生长, 使得其根系物质分配比显著高于生产种, 再一次影响了根茎的物质供给。

与原种、原原种相比, 生产种株高、茎粗增加, 分蘖减少, 为其高产奠定了理想的株型结构; 而物质积累速率高与快速增长期长, 使其物质积累显著增加, 为其高产奠定了物质基础。同时较低的茎鞘、叶片和根系物质分配比与较高的根茎物质分配比充分发挥其物质基础, 使得生产种的产量显著高于原种和原原种。较高的产量使得T₃总收入、效益和产投比均显著高于T₁和T₂, 表明生姜生产种较原种和原原种具有显著的产量优势, 能够有效提高生姜生产的经济效益。大量研究表明, 随组培世代的增加, 种块病原菌积累, 感病程度增加, 且产量和品质均大幅下降^{[21, 22, 23]}。综合考虑不同世代组培生姜产量差异与种源种性退化规律, 可以把生姜生产种(T₃)作为生产上的最佳推广世代, 既能充分发挥其产量潜力, 又能控制病害的发生。

(责任编辑侯春晓)

参考文献

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[12]	王林生, 孔祥生, 李友军, 等. 徐薯18脱毒薯苗不同世代干物质积累与分配规律研究[J]. 华中农业大学学报, 2000, 19(2): 99-101. WANG L S, KONG X S, LI Y J, et al. The law of accumulation and distribution of dry matter on four generation virus free cutting of Xushu 18[J]. Journal of Huazhong Agricultural, 2000, 19(2): 99-101. (in Chinese with English abstract) [本文引用:3]
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2017

0.0

... 生姜(Zingier officinal Roscoe)为姜科多年生草本植物,具有食用和药用双重价值,在世界范围内广泛种植^[1,2] ...

2018

0.0

... 生姜(Zingier officinal Roscoe)为姜科多年生草本植物,具有食用和药用双重价值,在世界范围内广泛种植^[1,2] ...

2015

0.0

... 我国生姜在全球市场占主导地位,栽培面积与总产量均居世界首位^[3] ...

2016

0.0

... 同时,生姜作为我国优势特色蔬菜和主要调味品原料,已成为乡村振兴与脱贫攻坚的特色高效产业,也是重庆市现代特色效益农业调味品产业体系主导作物之一^[4,5] ...

2014

0.0

李秀, 徐坤, 巩彪. 生姜种质遗传多样性和亲缘关系的SRAP分析[J]. 中国农业科学, 2014, 47(4): 718-726.

, XU

, GONG

. Genetic diversity and phylogenetic relationship of ginger germplasm resources revealed by SRAPs[J]. Scientia Agricultura Sinica, 2014, 47(4): 718-726. (in Chinese with English abstract)

【Objective】Ginger is vegetatively asexual crop that owns lots of local varieties, and its biological characteristics have been verified in many aspects caused by climate change, natural and artificial selection for thousands of years. This study was conducted to make a scientific classification of ginger germplasm, provide evidence for germplasm collection, protection and innovation of ginger by investigating the genetic diversity and relationship.【Method】Good genomic DNA was extracted from young leaves of 51 ginger accessions from different areas of the world following the CTAB method, and then were amplified by sequence-related amplified polymorphism molecular markers to analyze genetic diversity and phylogenetic relationship. Separation of the amplified fragments was performed on 6% denaturing polyacrylamide gels, The gels were stained with AgNO3 for visualizing the SRAP fragments, and then“0, 1” matrix was obtained according to the electrophoresis results. The number of polymorphic loci, percentage of polymorphic loci, effective number of alleles, Nei’s genetic similarity coefficient, genetic distance and the indexes of Nei’s genetic diversity and Shannon information were estimated by POPGENE version1.32. The cluster analysis of 51 ginger accessions based on unweighted pair-group method and 7 ginger populations based on Nei’s genetic distance were performed on NTSYS version2.10e to classify the ginger germplasm. Meanwhile, the origin and transmission of ginger were discussed in accordance with genetic diversity and phylogenetic relationship of ginger germplasm in different ecotopes and in light of the relevant characteristics of origin center and historical records.【Result】Among the 305 bands detected by 15 selective primer pairs, 188 (61.68%) were polymorphic. On average, each primer combination amplified 20.33 loci and 12.53 polymorphic loci. This showed that genetic variation of ginger is extensive. The average indexes of Nei’s genetic diversity and Shannon information were 0.3689 and 0.5510, respectively. Ginger germplasms were divided into 3 groups and 9 subgroups accordingly. Through comparative analysis, it was found that the same group gingers were from the same or similar region. Further analysis showed that ginger populations were separated into seven groups by geographical distribution, whose Shannon information index ranged from 0.2901 to 0.4807. The seven geographical populations were divided into 4 groups when the genetic similarity coefficient was set at 0.9. The Northern China population and Africa population formed their own group, respectively; Southeast Asia population, the Japan and Korea population formed a group; the coastal Southeastern China population, Southwest Plateau population and Central China population fell into another group.【Conclusion】 Analysis of 51 ginger accessions based on the indexes of Nei’s genetic diversity and Shannon information showed that genetic diversity of ginger was abundant though it is an asexual crop. The cluster analysis conducted with UPGMA of 51 ginger accessions based on genetic diversity coefficient indicated that genetic diversity of ginger germplasm was greatly influenced by geographical origins. Analysis on populations’ genetic diversity showed that genetic diversity of domestic populations in China was higher compared with foreign populations, in particular, Northern China population was far from other populations, moreover where the ginger cultivation has a long history, so the Northern China could be determined as the secondary center of origin; Africa is likely one of the native origin of gingers besides Southeast Asia, as the Africa population not only has a higher genetic diversity index, is far from other populations, and its space distribution is near from geometry center, but Africa also has wild species.

College of Horticulture Science and Engineering, Shandong Agricultural University /State Key Laboratory of Crop Biology/Key Laboratory of Biology and Genetic Improvement of Horticultural Crops in Huanghuai Region, Ministry of Agriculture, Taian 271018, Shandong

【目的】生姜为无性繁殖作物，千百年来，在经历气候变迁、自然选择和人工选择过程中，其生物学性状呈现出多方面变异，造就了许多地方品种。因此，研究生姜种质资源的遗传多样性及亲缘关系，对生姜资源进行科学分类，为生姜种质的收集、保护和创新利用提供理论依据。【方法】采用CTAB法提取生姜幼叶基因组DNA，利用SRAP（相关序列扩增多态性，sequence-related amplified polymorphism）分子标记技术，对来源于世界不同地区的51份生姜种质的基因组DNA进行多态性扩增，PCR产物采用6%变性聚丙烯酰胺凝胶电泳分离，采用银染法显色。根据电泳结果得到“0，1”矩阵，利用POPGENE version1.32软件计算每对引物的多态性位点数、多态性位点百分率，以及有效等位基因数Ne、Nei’s遗传相似系数（GS）、遗传距离（GD）、Nei’s遗传多样性指数、Shannon信息指数等指标，并采用NTSYS version2.10e软件进行种质间的UPGMA（非加权组平均法，unweighted pair-group method）聚类分析和基于Nei’s遗传距离的组群间的聚类分析，对生姜资源进行分类；同时，根据不同生态区之间生姜种质资源的遗传多样性和亲缘关系，结合植物起源中心相关特性及历史记载，探讨生姜的起源与传播。【结果】筛选出的15对引物共扩增出305条带，其中188条为多态性条带，多态性比率为61.68%，平均每对引物扩增出20.33个位点和12.53个多态性位点，说明生姜的遗传变异较为广泛。51份生姜种质的Nei’s遗传多样性指数、Shannon信息指数平均值分别为0.3689、0.5510，据此可将生姜种质分为3个大类、9个亚类，分析比较发现，每一类的生姜多来源于相同或相近区域。进一步研究表明，7个不同地理来源群体的Shannon信息指数范围是0.2901—0.4807，在遗传相似系数为0.9时，可将7个群体分为4类，华北群体、非洲群体各为一类，东南亚群体与日韩群体为一类，东南沿海群体、西南高原群体、华中群体为一类。【结论】品种间基于Nei’s遗传多样性指数和Shannon信息指数的分析表明，生姜虽为无性繁殖作物，但其遗传多样性较为丰富；根据遗传相似系数的UPGMA聚类分析结果，生姜种质资源的遗传多样性受地理来源影响较大；群体间遗传多样性分析表明，中国国内群体的遗传多样性高于国外群体，尤其华北群体与其他群体的遗传距离较远，且其栽培历史悠久，可以判定中国华北地区是生姜的次生起源地；而鉴于非洲群体的遗传多样性指数较高，与其他群体的遗传距离较远，且空间分布距中心位置较近，又有野生种存在，据此推断非洲可能是除东南亚之外的另一生姜原生起源地。

... 高产型生姜的主要特征为茎秆粗壮、分蘖较少、叶片较厚、同化系统发达、根茎干物质含量较低^[5,20] ...

2017

0.0

... 生姜以地下根茎无性繁殖为主,连年自留种,种性退化,病原物积累,土传病害严重,使得生姜生长缓慢,品质下降,减产甚至绝收,严重制约生姜产业的持续健康发展^[6,7] ...

2017

0.0

2015

0.0

... 而组织培养是解决无性繁殖作物种性退化、病原物积累,提高抗病性和产量潜力的重要措施^[8,9,10,11] ...

1997

0.0

宋伯符, 王胜武, 谢开云, 等. 我国甘薯脱毒研究的现状及展望[J]. 中国农业科学, 1997, 30(6): 43-48.

SONG B

, WANG S

, XIE K

, et al. Status and prospects of sweetpotato viruses elimination in China[J]. Scientia Agricutura Sinica, 1997, 30(6): 43-48. (in Chinese with English abstract)

国际马铃薯中心驻京办事处，北京 100081 江苏徐州甘薯研究中心山东省农业科学院植保所内蒙古大学生物系中国农业科学院生物技术中心山东省农业科学院作物所

概述了我国甘薯脱毒有关研究的现状并展望了这一领域未来的发展趋势。调查表明我国大多数甘薯田块植株的病毒感染率达60%～70%，有的甚至达到90%。徐州甘薯研究中心保存的甘薯资源材料有80%受病毒感染。叶片带斑点和花叶是甘薯病毒病的主要病症，其次为卷叶和皱叶，再次为老叶羽状花叶锈斑和环斑。引起病毒病的病原主要是甘薯羽状斑驳病毒、甘薯潜隐病毒和褪绿斑驳病毒。1991～1995年5省55个试验点的试验结果表明，脱毒甘薯的增产效果年平均为35.22%，脱毒甘薯通常在生产中可使用2～3年，脱毒甘薯对黑斑病和黑痣病也有防治效果。脱毒甘薯在中国的推广面积1995年已达9万hm#+2,并期望在近年内有很大的发展。脱毒薯的利用结合育成新品种的推广将会达到更好的效果；而抗病毒品种的选育则是解决病毒问题的根本途径。

... 而组织培养是解决无性繁殖作物种性退化、病原物积累,提高抗病性和产量潜力的重要措施^[8,9,10,11] ...

2007

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葛胜娟. 生姜组培苗的培育及其生产应用[J]. 中国农学通报, 2007, 23(5): 75-78.

GE S

. Study on culture of tube-plantlets of ginger and its application in production[J]. Chinese Agricultural Science Bulletin, 2007, 23(5): 75-78. (in Chinese with English abstract)

Ginger was bred by tissue culture technique, which showed the medium MS+6-BA2.0mg/L+NAA0.1mg/L is best. The tube-plantlets were tested by hardening and transplanting, which showed the harden plantlets is best to raise transplanting survival rate. The optimal treatment are both stronger plantlets and hardening plantlets. The first generation ginger tuber growing out of the tube-plantlets is smaller and healthy, which is suitable to use as seed tuber. The second generation ginger tuber is larger and higher yield, which has good value to trade and economic benefit。

(Jiaxing Vocational Technical College, Jiaxing 314036)

利用组织培养技术，对生姜进行茎尖培养，筛选出适合丛生芽快速繁殖的培养基配方（MS+6-BA2.0mg/L+NAA0.1mg/L）；对生姜组培苗进行炼苗和移栽试验，结果炼苗提高移栽成活率的作用最大，壮苗加炼苗移栽成活率最高,可达100%；组培苗生产的一代姜，姜块小、姜体健康，适宜用作种姜；二代姜姜块大、产量高，商用价值与经济效益好。

... 而组织培养是解决无性繁殖作物种性退化、病原物积累,提高抗病性和产量潜力的重要措施^[8,9,10,11] ...

2017

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... 而组织培养是解决无性繁殖作物种性退化、病原物积累,提高抗病性和产量潜力的重要措施^[8,9,10,11] ...

... 干物质积累与分配是作物产量形成的基础,脱毒甘薯干物质的积累,在整个生育期内均明显高于对照,其分配规律为:前期以地上部茎叶为主,中后期地下部块根迅速增加,从而提高产量^[11,12] ...

2000

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... 王林生等^[12]研究表明,不同世代组培甘薯增产幅度差异明显,茎尖苗增产效果明显高于原原种苗、原种苗和良种苗 ...

... 而进入根茎膨大期后,物质积累从地上向地下转移,根茎物质分配比和根冠比逐渐提高,茎叶分配比则随之下降,与王林生等^[12]在甘薯上的研究结果一致 ...

2007

0.0

... 汤青林等^[13]通过测定不同世代组培芋苗叶片生理指标和产量,综合评价指出组培2代要优于组培1代和普通苗,是大面积推广种植的种源 ...

2017

0.0

... 组培技术在甘薯、马铃薯上已有大量研究应用^[14,15] ...

2014

0.0

... 组培技术在甘薯、马铃薯上已有大量研究应用^[14,15] ...

2013

0.0

... 利用Logistic方程对生姜物质生产进行曲线拟合,可以得出最大理论生物量(final theoretical biomass yield,M_D)、最大生长速率(maximum growth rate,V_mD)、平均生长速率(average growth rate,V_aD)、最大生长速率出现时间点(time of instantaneous maximum slope,T_0D)、生长加速点(time of growth rate acceleration,T_1D),生长减速点(time of growth rate deceleration,T_2D)和快速生长期(rapid growth rate period,T_2D-T_1D)^[16,17,18] ...

2005

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2009

0.0

2015

0.0

卢会翔, 唐道彬, 吴正丹, 等. 甘薯产量、品质及农艺性状的基因型与环境效应研究[J]. 中国生态农业学报, 2015, 23(9): 1158-1168.

LU H

, TANG D

, WU Z

, et al. Genotypic variation and environmental effects on yield, quality and agronomic traits of sweet potato[J]. Chinese Journal of Eco-Agriculture, 2015, 23(9): 1158-1168. (in Chinese with English abstract)

The development of new sweet potato varieties with high starch and anthocyanin contents in storage roots has been the key objective of sweet potato research. In this study, natural variations in five white-fleshed sweet potato (with white or pale yellow flesh color) varieties (lines) and five purple-fleshed sweet potato (with purple flesh color) varieties (lines) were analyzed for the effects of environments on quality traits of storage roots in 2012 and 2013. Analyses of variance and correlation were conducted for the two types of sweet potato accessions in six regions of Chongqing City. The results showed significant differences ( P P r ≥ 0.700) among fresh yield, starch yield and dry root yield of sweet potato. Starch content had extremely positive correlations with dry root yield and starch yield ( r ≥ 0.363). For white-fleshed sweet potato accessions, stem diameter had extremely significantly positive correlation with vine length ( r = 0.439), extremely significantly negative correlation with starch content (r = 0.433), and significantly negative correlation with starch yield ( r = 0.318). For purple-fleshed sweet potato accessions, commodity rate was significantly positively correlated with branch number, starch yield and dry root yield ( r = 0.345, 0.368 and 0.357, respectively). In addition, a significantly negative correlation ( r = 0.397) was noted between vine length and branch number. It was concluded that affecting factors of yield, quality and agronomic traits of different genotypic accessions of sweet potato were different. Individualized selection and breeding approaches should be adopted on the basis of sweet potato types and breeding targets. The results provided valuable theoretical reference for screening advantageous environmental factors for quality traits formation, improvement of breeding approaches, and cultivation techniques of sweet potato. This could facilitate the promotion of quality improvement of sweet potato.

选育具有优质淀粉或花青素性状的甘薯新品种是甘薯育种的重要目标。本研究利用5个白肉(薯肉颜色为白色或淡黄色)甘薯品种(系)和5个紫肉(薯肉颜色为紫色)甘薯品种(系)的自然变异, 于2012年和2013年对重庆市6个地点2年间甘薯淀粉和花青素等性状测定结果进行方差分析、变异分析和相关性分析, 以期对甘薯产量、品质和农艺性状的自然变异规律和环境影响因素进行探索。结果表明, 基因型效应和环境效应下甘薯产量性状(鲜薯产量、淀粉产量、薯干产量)和品质性状(干率、淀粉含量、花青素含量)差异均达到显著水平( P P 环境效应>年份效应, 白肉甘薯产量性状的环境效应>基因型效应>年份效应, 紫肉甘薯产量性状的环境效应、年份效应大于基因型效应。重庆中海拔地区(万州区或酉阳县)甘薯产量相对较高, 低海拔地区(合川区、北碚区)甘薯薯块熟食性较好, 并且薯块熟食适口性评分低海拔地区>中海拔地区>高海拔地区(黔江区或巫溪县)。甘薯农艺性状(茎粗、最长蔓长、基部分枝数)和产量性状平均变异系数较大(CV=0.30, 0.28), 品质性状(干率、淀粉含量、薯块熟食适口性评分)平均变异系数较小(CV=0.11), 但花青素含量平均变异系数较大(CV=0.28)。相关性分析显示, 甘薯鲜薯产量、淀粉产量、薯干产量之间均呈极显著正相关( r ≥0.700); 淀粉产量、薯干产量均与淀粉含量呈极显著正相关( r ≥0.363)。白肉甘薯茎粗与最长蔓长呈极显著正相关( r =0.439), 与淀粉含量呈极显著负相关( r =0.433), 与淀粉产量呈显著负相关( r =0.318)。紫肉甘薯商品薯率与基部分枝数、淀粉产量、薯干产量均呈显著正相关( r =0.345, 0.368, 0.357), 基部分枝数与最长蔓长呈显著负相关( r =0.397)。白肉甘薯和紫肉甘薯在产量、品质及农艺性状的影响因子上存在不同, 在品种选育时应针对不同甘薯类型和育种目标采取不同选育方案。本研究可为筛选有利于甘薯品质性状形成的环境因子、促进育种手段和栽培技术的改进、加快甘薯品质改良提供理论依据。

... 作物产量与植株形态特征密切相关,茎基部较粗、蔓较短、分枝数较少、薯块大小均匀的甘薯往往产量较高^[19] ...

2012

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李秀, 徐坤, 巩彪, 等. 生姜农艺性状与产量形成关系的多重分析[J]. 中国农业科学, 2012, 45(12): 2431-2437.

, XU

, GONG

, et al. Multiple analysis of relationship of agronomic traits and yield formation in ginger[J]. Scientia Agricultura Sinica, 2012, 45(12): 2431-2437. (in Chinese with English abstract)

【Objective】 The purpose of this study was to provide academic evidence for germplasm evaluation and innovation of ginger, and identify the primary indexes of yield by investigating the variation of agronomic traits of ginger.【Method】The correlation, path analysis, principal components analysis and multiple linear regression analysis were conducted, whilst integrated the growth status of ginger, and applied critical-path analysis, to identify the primary indexes of ginger yield. 【Result】The results showed that the variation coefficient of agronomic traits of ginger reached 11.4%-51.3%, and the yield had a extremely significant correlation with stem diameter, stem FW, plant height, leaf FW and root FW, but a significant or extremely significant negative correlation with rhizome DM content and branch number. Plant height, stem diameter, root FW, stem FW and rhizome DM content were named as determinative morphological factors of high-yielding varieties, and branch number, leaf number and ginger bulb number were named as determinative morphological factors of low-yielding varieties. The determination coefficient of four factors selected by multiple linear regression analysis to ginger yield model was 0.8194.【Conclusion】The major determining factors of ginger yield were rhizome DM content, leaf FW, stem diameter and stem FW. The high yield varieties had thickest stem, fewer branch number, thicker leaf, developed assimilation system, and lower rhizome DM content.

山东农业大学园艺科学与工程学院/作物生物学国家重点实验室/农业部黄淮地区园艺作物生物学与种质创制重点实验室，山东泰安271018

【目的】研究分析生姜植株农艺性状变异特征，确定构成生姜产量的主要因子，为生姜种质评价与创新提供理论依据。【方法】通过对生姜农艺性状进行相关分析、通径分析、主成分分析及多元线性回归分析，结合生姜生长发育特点，运用统筹分析，明确生姜产量形成的主要决定因子。【结果】生姜主要农艺性状变异系数达11.4%—51.3%，且植株茎粗（0.7934）、茎鲜重（0.7302）、株高（0.6024）、叶鲜重（0.5847）和根鲜重（0.5039）均与产量呈极显著正相关，根茎干物质含量（-0.6155）、分枝数（-0.3680）与产量呈（极）显著负相关；株高、茎粗、根鲜重、茎鲜重、根茎干物质含量是高产型品种形态决定因子，而分枝数、叶片数、姜球数则为低产型品种形态决定因子；多元线性回归分析入选的4个因子对生姜产量模型的决定系数达0.8194。【结论】生姜单株产量的主要决定因子依次为根茎干物质含量（-0.3984）、叶鲜重（0.2866）、茎粗（0.2711）及茎鲜重（0.2624），高产型生姜品种的主要特征为茎秆粗壮、分枝较少、叶片较厚、同化系统发达、根茎干物质含量较低。

... 高产型生姜的主要特征为茎秆粗壮、分蘖较少、叶片较厚、同化系统发达、根茎干物质含量较低^[5,20] ...

2011

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... 大量研究表明,随组培世代的增加,种块病原菌积累,感病程度增加,且产量和品质均大幅下降^[21,22,23] ...

2006

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... 大量研究表明,随组培世代的增加,种块病原菌积累,感病程度增加,且产量和品质均大幅下降^[21,22,23] ...

2003

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... 大量研究表明,随组培世代的增加,种块病原菌积累,感病程度增加,且产量和品质均大幅下降^[21,22,23] ...