Mechanism of root exudates regulating plant responses to phosphorus deficiency
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摘要:
磷是植物生长发育的必需营养元素，土壤中有效磷含量低是限制作物高产的主要因素。由于长期不当施肥，土壤中累积大量磷素，其中大部分磷是植物难以直接吸收的难溶性无机磷和有机磷。植物在长期进化过程中形成了一系列适应低磷胁迫的机制，其中根系分泌物参与土壤磷活化利用的机制一直是研究的热点问题。本文总结了近十年来关于低磷胁迫调控根系分泌物(有机酸和紫色酸性磷酸酶)合成和分泌的研究进展，对根系分泌物在根际微生态中的重要作用提出了展望，旨在阐明通过控制作物根系分泌物来提高作物磷效率的途径，为培育磷高效作物品种和优化磷肥的田间管理提供思路和奠定理论基础。
Abstract:Phosphorus (P) is an essential nutrient for plant growth and development. Low phosphate (Pi) availability in soil largely limits crop yield. Due to long-term improper P fertilizer application, a lot of P accumulates to form a huge P pool in soil. However, most P are insoluble inorganic P and organic P, and are difficult to be directly absorbed by plants. Plants have evolved a set of adaptive strategies to low P stress. Among them, the mechanism of root exudates participating in P acquisition and utilization has always been a hot issue. In this review, advances in low P stress regulating synthesis and exudation of root exudates (organic acid and purple acid phosphatases) were summarized. Furthermore, the vital functions of root exudates in rhizosphere ecological system are discussed to elucidate mechanism of P efficiency enhancement in crops through root exudate regulation, which would provide some clues and theoretical bases for development of high P efficiency cultivar and optimization of Pi fertilizer management in fields.
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Keywords:
- low phosphorus /
- root exudate /
- purple acid phosphatase /
- organic acid
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自两系不育系被发现以来，两系法杂交已在水稻生产上得到应用，并显示出广阔的应用前景[1]。两系不育系育性不稳定，育性敏感期受外界环境的严重制约，如果该时期遇到异常天气，可能导致繁种失败。已经推广应用的两系不育系起点温度由于温度漂变，制种风险增加[2-3]，使得两系不育系的生产推广受到严重制约。此外，配合力不够理想也是其推广受阻的重要原因之一[4]。配合力包括一般配合力和特殊配合力，一般配合力指一个自交系和品种或其他一系列其他自交系和品种所产生的杂种一代的产量平均值；特殊配合力指在某个特定的杂交组合中2个自交系杂交产生的杂种一代的产量表现。一般配合力是评价亲本优良特性的重要依据，可通过一般配合力了解某亲本在杂交后代中的平均表现，特殊配合力是特定杂交组合中基因通过显性、上位性作用及与环境互作使后代表现相关优良性状的潜在能力。研究亲本的配合力对水稻杂交育种具有重要的指导意义，通过配合力评价种质资源在育种中的作用，可以充分利用水稻杂种优势，促进杂交水稻的发展[5]。若某亲本产量性状的一般配合力高，杂交组合的特殊配合力也较高，表明该亲本具有广泛的适用性，易选育高产优质的杂交组合[6]。遗传力反映亲本性状遗传给子代的能力[7]，为了探究性状的遗传力，可以把全部基因型方差占表现型方差的百分比作为广义遗传力(hB2)，把加性方差占表现型方差的百分比作为狭义遗传力(hN2)，用狭义遗传力度量性状的遗传力更可靠[8]。本研究对大穗型两系不育系‘M20S’主要穗部性状的配合力和遗传力进行研究，从生产实践出发，选用生产上广泛应用的7个优良杂交稻亲本进行不完全双列杂交(Incomplete diallel cross，NCⅡ)设计组配[9]，通过一般配合力、特殊配合力及遗传力分析，明确该不育系和恢复系在穗部性状上配合力的强弱，为优质高产杂交稻组合的选配提供参考依据。
1. 材料与方法
1.1 供试材料
光温敏核不育系：‘望S’、‘深08S’、‘Y58S’以及华南农业大学国家植物航天育种工程技术研究中心新选育的‘M20S’；恢复系：‘航恢1173’、‘航恢91’和‘航恢24’；4个不育系和3个恢复系配制的12个杂交组合，共计19份材料。
1.2 试验方法
试验在华南农业大学国家植物航天育种工程技术研究中心水稻育种试验田(N23°，E113°)进行。2017年早季以4个光温敏核不育系为母本和3个恢复系为父本，按照NCⅡ设计配制12个杂交组合；2017年晚季种植F1代，7月22日播种，8月7日水稻幼苗长到四叶一心时插秧，完全随机区组设计，3次重复，每个小区按照6×6规格种植，共36株，单本种植，田间管理措施与常规大田生产管理相同。完熟期时，从每个小区中选取3株有代表性的单株，用烘干机于45 ℃条件下干燥处理24 h，干燥后用量程40 cm的直尺测量穗长，用水稻数字化考种机YTS-5D考种并记录总粒数、结实率、千粒质量、单穗质量、一次枝梗数和着粒密度(每10 cm稻穗着生的水稻籽粒总粒数)。
1.3 数据与处理
数据分析采用SPSS 19.0和Microsoft Excel 2007进行，统计分析参照文献[10]的方法进行，配合力和遗传力分析按照文献[11-12]进行。根据固定模型估算试验材料的配合力效应，根据随机模型估算群体配合力方差和遗传参数。
2. 结果与分析
2.1 不同杂交组合F1代穗部性状的比较分析
考察各杂交组合F1代的穗部性状，统计分析各性状的平均值，结果见表1。‘M20S’配制的组合与‘望S’配制的组合相比，一次枝梗数、总粒数、单穗质量和着粒密度呈正向优势；与‘深08S’配制的组合相比，穗长、一次枝梗数、总粒数和着粒密度呈正向优势；与‘Y58S’配制的组合相比，一次枝梗数、总粒数、结实率、单穗质量和着粒密度基本呈正向优势。
表 1 12个杂交组合F1代穗部性状表型值Table 1. Phenotypic values of panicle traits in F1 generations of 12 hybrid combinations杂交组合
Hybrid combination穗长/cm
Panicle
length一次枝梗数
Primary branch number总粒数
Total grain number结实率/%
Seed setting rate单穗质量/g
Single panicle weight千粒质量/g
1 000-grain weight着粒密度
Grain
density望 S/航恢 1173
Wang S/Hanghui 117329.54 16.67 2 009.00 0.76 29.60 18.02 67.90 望 S/航恢 91
Wang S/Hanghui 9127.50 12.00 1 437.33 0.85 30.31 24.21 52.30 望 S/航恢 24
Wang S/Hanghui 2429.17 13.33 2 459.33 0.88 48.31 23.44 84.56 平均值 Mean value 28.74 14.00 1 968.56 0.83 36.08 21.89 68.25 深 08S/航恢 1173
Deep 08S/Hanghui 117327.74 14.00 2 352.33 0.81 38.97 18.59 85.01 深 08S/航恢 91
Deep 08S/Hanghui 9126.94 12.33 2 134.67 0.86 47.33 23.90 79.34 深 08S/航恢 24
Deep 08S/Hanghui 2426.67 12.67 1 908.00 0.91 37.92 23.71 71.73 平均值 Mean value 27.12 13.00 2 131.67 0.86 41.41 22.07 78.69 Y58S/航恢 1173
Y58S/Hanghui 117329.67 19.00 2 573.00 0.78 38.60 14.38 86.69 Y58S/航恢 91
Y58S/Hanghui 9128.81 11.00 1 256.67 0.76 21.95 22.56 43.69 Y58S/航恢 24
Y58S/Hanghui 2426.84 11.33 1 442.00 0.81 27.25 23.01 53.62 平均值 Mean value 28.44 13.78 1 757.22 0.78 29.27 19.98 61.33 M20S/航恢 1173
M20S/Hanghui 117330.36 17.00 2 382.33 0.89 24.62 19.40 78.49 M20S/航恢 91
M20S/Hanghui 9127.67 18.00 3 810.00 0.78 35.69 11.82 137.57 M20S/航恢 24
M20S/Hanghui 2425.56 16.00 3 581.00 0.79 54.74 18.98 141.00 平均值 Mean value 27.86 17.00 3 257.78 0.82 38.35 16.73 119.02 2.2 穗部性状配合力方差分析
7个穗部性状的配合力方差分析结果如表2所示，7个性状区间差异均不显著，组间差异均达极显著水平，说明不同杂交组合的基因型效应间存在真实的遗传差异。不育系母本中，穗长的一般配合力方差差异显著，一次枝梗数等其他6个性状的一般配合力方差差异极显著；恢复性父本中，总粒数和着粒密度的一般配合力方差差异显著，穗长等其他5个性状的一般配合力方差差异极显著；母本/父本组合中，穗长的特殊配合力方差差异显著，其他6个性状的特殊配合力方差差异极显著。表明杂交组合中7个性状均同时受亲本的一般配合力和杂交组合的特殊配合力的影响，即受基因的加性效应和非加性效应共同影响。
表 2 穗部性状配合力方差分析1)Table 2. Variance analysis of panicle trait combining ability方差来源
Source of variation穗长
Panicle
length一次枝梗数
Primary branch number总粒数
Total grain number结实率
Seed setting rate单穗质量
Single panicle weight千粒质量
1 000-grain weight着粒密度
Grain
density区间 Interplot 1.45 3.11 6 140.11 0.00 11.18 0.02 32.96 组间 Intergroup 6.38** 22.87** 823.60** 0.01** 307.15** 48.99** 2 750.49** 母本 Female parent 4.61* 27.78** 4 045 022.10** 0.01** 239.36** 55.19** 5 991.79** 父本 Male parent 16.33** 44.45** 128 764.19* 0.01** 303.20** 67.79** 318.64* 母本/父本 Female/Male 3.95* 13.22** 1 364 410.82** 0.01** 342.36** 39.61** 1 940.45** 误差 Error 1.36 1.96 36 785.08 0.00 11.03 0.98 67.00 1)“*”和“**”分别表示达 0.05 和 0.01 显著水平
1) “*” and “**” indicated significance at 0.05 and 0.01 levels, respectively2.3 穗部性状一般配合力和特殊配合力效应分析
4个不育系和3个恢复系亲本的7个性状的一般配合力分析结果如表3所示。相同性状不同亲本和不同性状相同亲本材料间的一般配合力效应不同，表明不同亲本不同性状的遗传基因效应复杂。
表 3 穗部性状一般配合力效应值Table 3. The effect value of general combining ability of panicle trait% 亲本
Parent穗长
Panicle
length一次枝梗数
Primary branch number总粒数
Total grain number结实率
Seed setting
rate单穗质量
Single panicle weight千粒质量
1 000-grain weight着粒密度
Grain
density望 S Wang S 2.49 −3.08 −13.61 0.67 −0.55 8.54 −16.54 深 08S Deep 08S −3.30 −10.00 −6.46 4.58 14.15 9.41 −3.89 Y58S 1.44 −4.62 −22.89 −4.99 −19.32 −0.91 −24.92 M20S −0.63 17.69 42.96 −0.27 5.72 −17.03 45.35 航恢 1173 Hanghui 1173 4.60 15.38 2.21 −1.75 −9.18 −12.75 −2.71 航恢 91 Hanghui 91 −1.10 −7.69 −5.23 −1.15 −6.76 2.25 −4.29 航恢 24 Hanghui 24 −3.50 −7.69 3.02 2.90 15.94 10.50 7.00 ‘M20S’在一次枝梗数、总粒数和着粒密度性状上一般配合力最佳，明显高于其他不育系，单穗质量一般配合力表现为正值，穗长、结实率和千粒质量表现为负值，一般配合力好的性状较多，表明该不育系能通过提高一次枝梗数和着粒密度来提高总粒数，从而提高库容量，与优势互补的恢复系进行配组，易选育出产量潜力高的品种。在3个恢复系中，‘航恢24’在总粒数、结实率、单穗质量、千粒质量和着粒密度性状上一般配合力具佳，优势比较明显，可以与‘M20S’优势互补。
不同杂交组合的7个性状的特殊配合力分析结果如表4所示，相同性状不同组合间及相同组合不同性状间的特殊配合力效应值存在明显差异，表明基因互作具多样性。从单穗质量上看，‘Y58S’/‘航恢1173’特殊配合力效应值最高，‘深08S’/‘航恢24’最低，特殊配合力效应值的变幅在–25.54~34.89之间。从经济学产量相关性状上看，‘望S’/‘航恢24’、‘深08S’/‘航恢91’、‘Y58S’/‘航恢1173’、和‘M20S’/‘航恢24’的特殊配合力效应较好；‘M20S’配制的3个组合中，‘M20S’/‘航恢24’一次枝梗数、总粒数、单穗质量、千粒质量和着粒密度这5个经济性状的特殊配合力表现为正效应，特别是总粒数、单穗质量和着粒密度这3个性状的特殊配合力效应值较高，该杂交组合在以‘M20S’为母本的3个组合中最符合大穗型育种的要求。
表 4 穗部性状特殊配合力的效应值Table 4. The effect value of special combining ability of panicle trait% 杂交组合
Hybrid combination穗长
Panicle
length一次枝梗数
Primary branch number总粒数
Total grain number结实率
Seed setting rate单穗质量
Single panicle weight千粒质量
1 000-grain weight着粒密度
Grain
density望 S/航恢 1173
Wang S/Hanghui 1173−1.72 3.08 −0.44 −6.74 −8.67 −6.44 2.47 望 S/航恢 91
Wang S/Hanghui 91−3.30 −6.15 −18.08 3.98 −9.13 9.24 −15.21 望 S/航恢 24
Wang S/Hanghui 245.03 3.08 18.52 2.76 17.80 −2.80 12.74 深 08S/航恢 1173
Deep 08S/Hanghui 1173−2.38 −8.46 7.47 −4.58 2.45 −4.47 10.39 深 08S/航恢 91
Deep 08S/Hanghui 910.47 3.08 5.36 1.28 23.09 6.83 5.16 深 08S/航恢 24
Deep 08S/Hanghui 241.91 5.38 −12.83 3.30 −25.54 −2.36 −15.55 Y58S/航恢 1173
Y58S/Hanghui 1173−0.20 20.77 33.59 0.94 34.89 −15.06 33.74 Y58S/航恢 91
Y58S/Hanghui 912.42 −11.54 −16.74 −1.68 −13.40 10.54 −17.41 Y58S/航恢 24
Y58S/Hanghui 24−2.22 −9.23 −16.85 0.74 −21.49 4.52 −16.33 M20S/航恢 1173
M20S/Hanghui 11734.31 −15.38 −40.63 10.38 −23.68 25.97 −46.60 M20S/航恢 91
M20S/Hanghui 910.42 14.62 29.46 −3.57 −5.56 −26.61 27.46 M20S/航恢 24
M20S/Hanghui 24−4.72 0.77 11.17 −6.81 29.24 0.64 19.14 此外，对亲本一般配合力效应和杂交组合特殊配合力效应进行比较，发现亲本一般配合力效应与杂交组合特殊配合力效应似乎是相对独立的，亲本一般配合力高的，杂交组合特殊配合力不一定高，亲本一般配合力低的，杂交组合特殊配合力不一定低。
2.4 穗部性状配合力的基因型方差估算
估算穗部各性状的一般配合力和特殊配合力基因型方差，可以更深入地了解双亲及其互作对杂种后代性状的影响，估算结果见表5，通过σ122与σ12+σ22以及Vg与Vs对比可知，总粒数、结实率、千粒质量、着粒密度和单穗质量的σ1-22>σ12+σ22，且Vs>Vg，表明这些性状以受亲本互作非加性效应的影响为主。穗长和一次枝梗数的σ1-22<σ12+σ22、Vs<Vg，表明这2个性状以受亲本基因加性效应影响为主。通过σe2与σG2对比可知，所有性状的σG2>σe2，表明亲本各性状受遗传的影响为主，受环境影响占次要地位，F1的各个性状受遗传与环境共同影响。
表 5 穗部性状配合力的基因型方差及贡献率1)Table 5. Genotypic variance and contribution rate of combining ability of panicle trait性状 Trait σ12 σ22 σ1-22 σe2 σ12+σ22 穗长 Panicle length 0.055 0 1.375 6 0.861 7 1.364 9 1.430 6 一次枝梗数 Primary branch number 1.213 0 3.469 1 3.754 2 2.477 3 4.682 1 总粒数 Total grain number 223 384.270 0 −137 294.100 0 442 541.910 0 36 785.081 0 86 090.203 0 结实率 Seed setting rate 0 −0.000 5 0.002 1 0.002 8 −0.000 4 单穗质量 Single panicle weight −8.583 3 −4.351 1 110.443 5 11.029 6 −12.934 4 千粒质量 1 000-grain weight 337.611 7 −180.201 1 624.485 0 0.978 2 157.410 6 着粒密度 Grain density 1.298 3 3.131 1 12.877 3 0.978 2 4.429 4 性状 Trait σG2 σP2 Vg/% Vs/% 穗长 Panicle length 2.292 3 3.657 1 62.41 37.59 一次枝梗数 Primary branch number 8.436 3 1 091.360 0 55.50 44.50 总粒数 Total grain number 528 632.120 0 565 417.200 0 16.29 83.71 结实率 Seed setting rate 0.001 7 0.004 6 −25.73 125.73 单穗质量 Single panicle weight 97.509 0 108.538 6 −13.26 113.26 千粒质量 1 000-grain weight 781.895 6 782.873 8 20.13 79.87 着粒密度 Grain density 17.306 7 18.284 9 25.59 74.41 1) σ12：P1（一套n1＝4的不育系亲本）的一般配合力基因型方差；σ22：P2（一套n2＝3的恢复系亲本）的一般配合力基因型方差；σ1-22：P1-2（亲本互作）的特殊配合力基因型方差，又叫显性方差；σe2：环境方差；σ12+σ22：一般配合力加性基因型方差；σG2：总基因型方差；σP2：表现型方差；Vg：一般配合力方差，反映加性效应；Vs：特殊配合力方差，反映非加性效应
1) σ12: P1 (a set of n1=4 male sterile parents) general gratification genotype variance; σ22: P2 (a set of n2=3 restorative parents) general gratification genotype variance; σ1-22: P1-2 (parent interaction) special combining ability genotype variance (also called dominant variance); σe2: environmental variance; σ12+σ22: General combining ability additive genotype variance: σG2: Total genotype variance; σP2: Phenotypic variance; Vg : General combining force variance; Vs: Special combining force variance, reflecting non-additive effect2.5 穗部性状的遗传力
7个穗部性状的遗传力如表6所示。广义遗传力从大到小依次为：千粒质量、着粒密度、总粒数、单穗质量、一次枝梗数、穗长和结实率。所有性状的广义遗传力均比较大，除了结实率广义遗传力为37.49%，其余性状的广义遗传力都在60%以上，其中千粒质量和总粒数的广义遗传力达90%以上，说明这些性状很大程度上受遗传效应的影响。狭义遗传力从大到小依次为：一次枝梗数、穗长、着粒密度、千粒质量、总粒数、结实率和单穗质量，这些性状的狭义遗传力都在45%以下，遗传稳定性一般，性状的遗传力较弱，特别是结实率和单穗质量的狭义遗传力均小于0，影响非常显著，后代遗传稳定性差，亲本性状容易与自然环境、栽培方式等因素互作，对组合性状表现有直接影响。
表 6 各性状遗传力的估算1)Table 6. Estimation of heritability of each trait% 性状 Trait hB2 hN2 穗长 Panicle length 62.68 39.12 一次枝梗数 Primary branch number 77.30 42.90 总粒数 Total grain number 93.49 15.23 结实率 Seed setting number 37.49 −9.65 单穗质量 Single panicle weight 89.84 −11.92 千粒质量 1 000-grain weight 99.88 20.11 着粒密度 Grain density 94.65 24.22 1) hB2：广义遗传力；hN2：狭义遗传力
1) hB2: Generalized heritability; hN2: Narrow heritability3. 讨论与结论
3.1 杂交水稻穗部性状的遗传特点
穗部性状的一般配合力和特殊配合力方差差异均达显著或极显著水平，说明这些性状的遗传是受加性效应和非加性效应共同控制的。这些性状的配合力方差分析结果表明一次枝梗数和穗长的一般配合力方差较大，说明这2个性状受加性效应的影响较大；总粒数、结实率、千粒质量、着粒密度以及单穗质量的特殊配合力方差较大，说明这些性状主要受非加性效应的影响。此外，对亲本一般配合力效应和杂交组合特殊配合力效应进行比较，发现亲本的一般配合力效应与杂交组合的特殊配合力效应似乎是相对独立的，与前人研究情况不完全相同[13-14]，亲本一般配合力高的，组合的特殊配合力不一定高，亲本一般配合力低的，组合的特殊配合力不一定低，与前人研究一致[15-17]。由穗部性状广义遗传力分析可知，总粒数、千粒质量、着粒密度和单穗质量表现突出，受遗传效应的作用极大。在优质杂交稻亲本的改良中，一次枝梗数、穗长等狭义遗传力高的性状，可在杂交早代选择，以提高育种效率。
3.2 大穗型不育系‘M20S’的综合利用评价
在亲本选配的过程中，需要综合考虑亲本的一般配合力与杂交组合的特殊配合力才能获得优良组合[18-19]，根据研究分析，‘M20S’在总粒数、一次枝梗数、着粒密度性状上一般配合力最突出，单穗质量上一般配合力也是正值，表现良好，该不育系是一个大穗型的不育系，而穗型的大小是通过总粒数来分类的，总粒数的一般配合力达到了42.96%，远远超过其他亲本，说明‘M20S’的大穗性状不但能通过杂交遗传给后代，而且该不育系可以通过提高一次枝梗数来提高总粒数，从而提高经济学产量，是一个优良的亲本。对于杂交组合‘M20S/航恢24’，总粒数、着粒密度和单穗质量的特殊配合力较高，其中单穗质量的特殊配合力较大，为29.24%，其他性状特殊配合力效应较好，表明‘M20S/航恢24’在‘M20S’组配的3个组合中是最符合大穗型育种要求的组合。
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图 1 植物紫色酸性磷酸酶进化树分析
Ac: 洋葱Allium cepa; As: 紫云英Astragalus sinicus; At: 拟南芥Arabidopsis thaliana; Ec: 轮花大戟Euphorbia characias; Gm: 大豆Glycine max; Hv: 大麦Hordeum vulgare; Ib: 甘薯Ipomoea batatas; La: 白花羽扇豆Lupinus albus; Ll: 黄花羽扇豆Lupinus luteus; Mt: 截形苜蓿Medicago truncatula; Nt: 烟草Nicotiana tabacum; Os: 水稻Oryza sativa; Pt: 枳Poncirus trifoliate; Pv: 菜豆Phaseolus vulgaris; St: 马铃薯Solanum tuberosum; Ta: 小麦Triticum aestivum; Zm: 玉米Zea mays
Figure 1. Phylogenetic tree analysis of plant purple acid phosphatase
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