P高效利用杉木无性系适应环境磷胁迫的机制研究
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摘要
杉木[Cunninghamia Lanceolata (Lamb.) Hook]是我国南方最重要的造林树种之一。随着杉木造林面积的不断扩大,阔叶林面积锐减,杉木越来越多地在同一林地上连续栽植,多代连栽导致林地生产力下降,出现了人工林经营的“第二代效应”,已严重影响了杉木人工林的持续经营。我国南方林区20世纪80 - 90年代营造的大面积杉木速生丰产林,多已进入主伐阶段,均面临二代更新问题。目前杉木人工林地力衰退防治措施主要是通过改良土壤条件来促进杉木生长,这种传统的改地适树防治方法较少考虑通过选择养分利用效率高的基因型来适应林地的土壤条件,特别是对在较低肥力水平的林地上能获得高产的营养高效利用基因型研究甚少,当前速生的杉木基因型对林地养分的利用效率不一定是最高,有的是以牺牲林地的养分为代价来获得高产,这种基因型虽然速生,但对地力消耗也大,因此截止目前有关杉木人工林地力衰退降防治效果不明显,仍未研究出可供生产上大面积推广应用的防治杉木连栽生产力下降的有效措施,因此如何解决南方林区大面积杉木林退化地的造林更新成为当前林业生产中急需解决的重大课题。
本文针对南方林区土壤有效磷低但全磷丰富的特点,从提高杉木对林地磷素的利用效率入手,通过对我国杉木中心产区不同杉木无性系子代测定林生长和磷素利用特性的全面调查,进行不同杉木无性系子代测定林生长和P素利用效率的比较研究,在此基础上初步筛选出P高效利用杉木速生无性系。利用初选的12个P高效利用速生杉木无性系为研究对象,通过均一性磷胁迫模拟试验,分析不同杉木无性系对环境低磷胁迫的形态学和生理学响应差异,探讨P高效利用杉木无性系适应低磷胁迫内在机制,进一步筛选对土壤难溶性磷活化能力高和耐低磷能强的P高效利用杉木无性系。最后利用对土壤难溶性磷活化能力高和耐低磷能强的2个P高效利用杉木无性系进行异质供磷胁迫模拟试验,通过设计不同钙梯度低磷水培处理,通过异质供磷胁迫条件下杉木无性系形态学和生理学特异性指标的测定,探讨钙在P高效利用杉木无性系对环境低磷胁迫信号感受中的作用,分析P高效利用杉木无性系适应环境磷胁迫的内在机制,为揭示杉木耐低磷营养性状的生态遗传和完善林木营养遗传学方法提供科学依据。主要的研究结果如下:
1、7年生84个不同杉木无性系子代测定林的生长存在明显差异。不同无性系子代测定林树高、胸径和材积的年均生长量分别为0.84 m·a-1、1.12 cm·a-1、0.0036 m3·a-1。初步筛选出M8、M10、M11、M23、M28、M29、M30、M32、M33、M34、M101、M102、M136、M151、M211、M239和M249等17个速生杉木无性系,占参试无性系总数的19.77 %。
2、17年生45个不同杉木无性系子代测定林的生长存在明显差异。不同杉木无性系子代测定林树高、胸径和材积的年均生长量分别为0.94 m·a-1、1.22 cm·a-1、0.0161 m3·a-1。初步筛选出了M17、M34、M35、M40和M43号等5个速生无性系,占参试无性系总数的11.11 %。与初级嫁接种子园实生苗相比,17个无性系树高、胸径和材积平均现实增益值分别为11.81 %、15.50 %和37.16 %。
3、17年生不同杉木无性系的全树P利用效率平均为4886.67 g·g-1。其中M1利用效率最高,是不同无性系平均值的1.27倍,依次排序为M17>M10>M9>M45>M23>M41>M3>M19>M24>M43>M8>M25>M37>M34。不同无性系干材P利用效率大小排序为M1>M10>M9>M17>M41>M23>M19>M45>M3>M43>M8>M25>M24>M37>M34,其中M1比平均值增加25.84 %,是M34的1.57倍。
4、初选21个P高效利用杉木速生无性系在不同P处理条件下根系生长差异明显。难溶性Fe-P充足处理中,不同杉木无性系根系长度、表面积、平均直径和体积的平均值分别为15.58 m、566.56 cm2、0.77 mm和17.13 cm3,其中M8、M9、M11、M39和M302的根系发育较好,说明这7个杉木无性系在土壤有效态磷匮乏但难溶性磷丰富的逆境中根系觅养能力较强。缺磷处理中M1、M3、M9、M17、M21和M302根系发育较好,可见这4个杉木无性系耐磷饥饿能力较强。
5、初选21个P高效利用杉木速生无性系在不同P处理条件下生物量差异较大。与正常供磷相比,难溶性磷充足处理条件下生物量平均增加24.06 %,其中M302、M9、M8、M1、M11、M17、M39和M19增幅较大,说明它们在有效磷缺乏但难溶性铁磷充足逆境中觅养能力较强,在体内抗逆机制调节作用下生物量反而高于正常供磷。缺磷处理中,不同杉木无性系生物量平均减小31.97 %,其中M39、M1和M302耐磷饥饿能力较强,保持较高生产力,这可能与其体内磷素利用效率较高有关。
6、不同P处理对初选21个P高效利用杉木无性系体内P素积累与分布影响不同。参试21个不同杉木无性系各器官P素含量平均值的高低均表现为:叶>根>茎。其中根P素含量受低磷胁迫影响显著,这可能是杉木无性系遭遇低磷胁迫时,一方面体内P素向地下部转移,提高根系生产力,以增强对土壤P素的觅养能力;另一方面是由于根系活化了土壤中难溶性磷而使根的P素含量增多。铁磷处理中,M5、M17、M19、M21、M24、M37、M41、M48和M302号杉木无性系根P素含量较高;缺磷处理中,M3、M4、M5、M19、M21、M24、M28、M39、M102和M302根P素含量较高。
7、不同P处理对不同杉木无性系P素利用效率的影响存在差异。在正常供磷、铁磷处理和缺磷处理中,参试21个杉木无性系PUE的平均值分别为1.826 kg·g-1、1.663 kg·g-1和1.814 kg·g-1。铁磷处理条件下M3、M4、M19和M38号杉木无性系PUE比正常供磷处理大,增幅为2.48 %～28.04 %;缺磷处理条件下M41、M1、M19、M9、M17、M11、M37、M24、M21和M211均比正常供磷处理大。
8、随着低磷胁迫时间的延长,参试21个不同杉木无性系叶绿素总量的平均值表现出先快速增加,再缓慢增加的趋势;铁磷处理表现出同样变化规律。缺磷处理中,叶绿素总量先迅速增加,后增加速度变缓,最后减小。铁磷处理的胁迫末期,M3、M18、M24和M102号杉木无性系叶绿素总量的增幅比正常供磷处理大。缺磷处理的胁迫末期仅有M5、M18和M41等3个杉木无性系叶绿素总量增幅比正常供磷处理大。
9、低磷胁迫越严重,杉木叶片MDA含量升高越快。随着试验时间的延长,正常供磷处理中参试21个不同杉木无性系叶片MDA含量的平均值表现为先缓慢下降,再较大幅度下降,最后缓慢增加的变化趋势;铁磷处理变化规律与正常供磷处理相似;缺磷处理表现出不同变化规律:先缓慢上升,再急剧下降,最后缓慢增加。与正常供磷处理的胁迫末期相比,铁磷处理中M4、M19、M39、M102、M211和M302叶片MDA含量增幅较小,;缺磷处理中是M4、M5、M11、M18、M37、M38、M48和M211叶片MDA含量增幅较小。
10、随着试验时间的延长,不同P处理中参试21个杉木无性系叶片可溶性蛋白含量的变化表现出相似的规律:升高→下降→缓慢下降至胁迫前水平。但不同杉木无性系叶片可溶性蛋白含量的变化规律差异悬殊。与正常供磷处理中杉木无性系叶片可溶性蛋白含量的增幅相比,铁磷处理胁迫初期,M1、M3、M4、M8、M18、M19、M24、M28、M37、M38、M48和M211号的增幅较大,表明这12个无性系对低磷胁迫较敏感,可能是由于它们能够迅速启动体内酶等抗逆蛋白活性物质的生物合成;至胁迫末期,M11、M18、M19、M37、M38和M211的增幅较大,可能是铁磷处理逆境条件下这6个杉木活化难溶性铁磷能力较强,有效缓解了体内正常蛋白质合成受阻困境,且抗逆蛋白合成能力较高。缺磷处理低磷胁迫初期时,M1、M3、M4、M8、M9、M11、M19、M21、M24、M28、M38、M39、M41、M48和M211号的增幅较大;至胁迫末期,M3、M4、M11、M18、M19、M21、M24、M38、M39和M211号的增幅较大。
11、随着试验时间的延长,铁磷处理和缺磷处理条件下初选21个P高效利用杉木速生无性系叶片APA活性的平均值的变化规律与正常供磷处理相似:缓慢下降→升高→下降至较低水平的变化规律。然而,不同杉木无性系叶片APA活性的变化规律差异显著。铁磷处理胁迫初期, M1、M8、M9、M11、M17、M19、M39、M48和M211叶片APA活性增幅比正常供磷处理大;至胁迫末期,M3、M5、M8、M9、M11、M18、M19、M24、M28、M37、M38、M39和M211比正常供磷处理大。缺磷处理低磷胁迫初期,M1、M8、M38、M41和M211叶片APA活性增幅比正常供磷处理大;胁迫末期M1、M3、M5、M8、M18、M19、M24、M28、M37、M39、M102、M211和M302比正常供磷处理大。
12、初选21个P高效利用杉木速生无性系在不同P处理条件下干物质积累与各生理学指标之间相关性不强,但与不同形态学指标密切相关。铁磷充足处理中,根长、根表面积、根直径和根体积的增量与杉木无性系干物质积累量的相关系数分别达0.640、0.686、0.682和0.658,P值检验均呈极显著相关;缺磷处理条件下,根长、根面积和根体积对干物质产量影响显著,苗高也是决定干物质积累的相关因子之一。PUE与不同形态学指标相关性不大,不同生理学指标中叶绿素a和叶绿素总量均与杉木无性系PUE呈显著负相关。铁磷处理中,杉无性系PUE与叶片可溶性蛋白含量和APA活性分别呈显著水平和极显著水平,说明在林地有效磷匮乏但全磷丰富条件下,叶片的可溶性蛋白含量及APA活性可以作为评价筛选P素高效利用杉木无性系的可靠生理指标。
13、参试21个杉木无性系中M39号杉木无性系在有效磷匮乏全磷丰富的环境中适应性较强,且能保持较高的生物量积累速度,对P素的利用效率也较高,属于P高效利用杉木速生基因型;而M3、M4、M18、M19和M38的PUE较高但生物量产量较低,M1、M8、M9、M11、M17和M302具速生丰产特性,但PUE较低,可见这11个杉木无性系的开发潜力大,建议加强改良研究。缺磷处理试验表明:M1号杉木无性系耐磷饥饿特性明显,在极度缺磷的逆境中,能维持较高的根生长和干物质积累速度,可作为杉木营养遗传学研究的首选材料;M9号杉木无性系在完全缺P逆境中PUE较高,根生长速度较快,但生物积累量较小,这可能是与其通过扩大根系觅养能力较强,但体内P素的转移重复利用的能力较差有关。
14、不同异质供磷处理对M1和M4号杉木无性系根系生长的影响差异明显。两室均为正常磷供处理中,M1和M4两室根系生长差异均不大。高浓度铁磷、低浓度铁磷和缺磷处理条件下,M1和M4室Ⅰ中的根系长度、表面积、平均直径和体积生长量均比室Ⅱ大,但随着生长介质中磷素全量的减少,M1根系长度、表面积、平均直径和体积生长量均呈一定增加的趋势;而M4在生长介质中含有难溶性铁磷时根系生长速度较完全缺P时慢,这可能与它活化铁磷能力较强有关,一旦体内磷素充足,其通过根系延伸进行觅养的能力减弱。
15、不同异质P处理对M1和M4号杉木无性系体内P素分布的影响不同。M1号杉木无性系正常供磷处理条件下根与茎的P素含量相差最小,而在缺磷处理中差异最为明显;随着低磷胁迫加重,根与叶P素含量的差值递减趋势。通过对M1号杉木无性系室Ⅰ和室Ⅱ中根P素含量的比较,高浓度铁磷、低浓度铁磷和缺磷处理条件下室Ⅰ的根P素含量分别比室Ⅱ大0.18g·kg~(-1)、0.11g·kg~(-1)和0.35g·kg~(-1),表明M1号杉木在土壤磷极度匮乏逆境中耐磷饥饿杉木无性系可通过根系磷素分配格局的变化来维持根系的正常生长。
M4号杉木无性系正常供磷、高浓度铁磷、低浓度铁磷和缺磷处理根系平均P素含量分别比叶小了0.36 g·kg~(-1)、0.12 g·kg~(-1)、0.48 g·kg~(-1)和0.10 g·kg~(-1)。根与叶P素含量相比,不同P处理中缺磷处理中两者差异最小,而低浓度铁磷处理中两者差异较大。通过对M4号杉木无性系室Ⅰ和室Ⅱ中根P素含量的比较,高浓度铁磷、低浓度铁磷和缺磷处理条件下室Ⅰ中根的P素含量分别比室Ⅱ大0.11 g·kg~(-1)、0.01 g·kg~(-1)和0.04 g·kg~(-1),表明高浓度铁磷处理中两室根系P含量差异最大,说明在土壤有效磷匮乏但全磷丰富逆境中,活化铁磷能力较强杉木无性系根系磷素含量较大。
16、不同异质供磷处理对M1号和M4号杉木无性系PUE的影响差异较大。随着铁磷胁迫加重,M1号杉木无性系PUE呈逐渐升高趋势,与正常供磷处理相比,高浓度铁磷处理和低浓度铁磷处理PUE的值分别减小了21.20 %和2.94 %,而P0处理增大了7.78 %,说明土壤P含量越少,M1体内养分回收利用效率越高,耐磷饥饿越强;M4号杉木无性系PUE呈缓慢升高趋势,缺磷处理中PUE达2.41 kg·g~(-1),分别比正常供磷处理、高浓度铁磷和低浓度铁磷处理大了12.88 %、6.54 %和4.64 %。通过不同P处理M1和M4号杉木无性系PUE的比较,M4号杉木无性系PUE均比M1大,这可能与它们对铁磷活化能力不同有关。
17、不同异质供P处理对M1和M4号杉木无性系各生理指标的影响差异明显。不同异质供P处理对M1号杉木无性系各生理指标影响的差异表现为:正常供磷处理叶绿素总量表现出升高→缓慢降低→升高的变化规律;高浓度铁磷处理和缺磷处理呈缓慢升高→缓慢降低的变化趋势。叶片MDA含量变化规律的差异:正常供磷和缺磷处理条件下呈升高→降低的变化趋势;高浓度铁磷处理缓慢增加,而低浓度铁磷处理表现出升高→降低→缓慢升高的变化规律。叶片可溶性蛋白含量变化规律的差异:不同处理均随着胁迫时间的延长,表现出先快速升高,再逐渐降低的变化趋势。叶片APA活性变化规律的差异:不同P处理均表现为缓慢变化之后,快速增强,再减弱的变化趋势。
不同异质供P处理对M4号杉木无性系各生理指标影响的差异表现为:不同P处理叶绿素含量均表现出升高→降低→缓慢升高的变化规律,其中缺磷处理变化幅度很小。叶片MDA含量变化规律的差异:不同处理均表现出缓慢升高→降低的变化趋势,其中正常供磷处理降幅最大,缺磷处理最小。叶片可溶性蛋白含量变化规律的差异:不同处理均随着胁迫时间的延长,表现出先快速升高,再逐渐降低的变化趋势。叶片APA活性变化规律的差异:不同P处理均表现为缓慢变化之后,快速增强,再减弱的变化趋势。
18、不同Ca梯度在杉木感受低磷胁迫的试验表明:低磷胁迫条件下随胁迫时间的延长,水培杉木营养液酸度呈先上升后下降的变化规律。与无磷无钙处理相比,加钙处理有助于调节杉木根系生理活性,促进根系分泌酸性物质。随着Ca梯度的减小,杉木根P积累量所占百分数先依次降低再升高;叶和茎的变化规律不明显,但两者总是呈相反方向增减。不同Ca梯度中,处理Ⅰ的PUE最高,其余6个处理差异未达显著水平,其中对照最低,值为0.717 kg·g~(-1),表明高钙处理对低磷胁迫杉木PUE有一定的影响,但变化规律不明显,这可能与水培环境持续缺磷且试验培养期较短有关。不同Ca梯度处理条件下,低磷胁迫杉木各生理学指标的变化规律不明显。随着水培施Ca浓度降低,低磷胁迫杉木叶片MDA含量呈先下降后升高再降低的变化趋势,叶绿素总量的变化规律与MDA相近;叶片可溶性蛋白含量与APA活性的变化规律相似,总体趋势表现为降低→升高→降低。有关Ca和P的互作在杉木感受环境低磷胁迫中的作用还需进一步深入研究。
Chinese fir (Cunninghamia Lanceolata) is the unique important fast-growing timber species in southern China. With the growing planting area of Chinese fir, broadleaved forests area reduced sharply, more and more Chinese fir planted continuously at the same ground, which caused a decline in plantation productivity, so that the arisen "second generation effect" of plantation management had a serious impact on the continued operation of Chinese fir plantation. A large area of the fast growing Chinese fir forest has built during the 1960s and 1970s in southern China forests, most of which have entered into the final cutting stage and all faced with the problem of second-generation update. Thus settlement of soil degradation caused by second- generation update of large area Chinese fir plantation in the South has become the major subject,which starve for solution in the current forestry production. At present, prevention and control measures of soil degradation in Chinese fir plantation mainly promote the growth of Chinese fir through the improvement of soil conditions.This traditional control methods of changing land with trees are with little regard for the efficiency of nutrient use by selecting the high forest trees to adapt to the soil conditions to study, especially efficient use of nutrition genotypes are rarely researched to gain high yields at the lower level of fertility in forest land. The current fast-growing genotypes of forest trees is not necessarily the highest in nutrient use efficiency, some get high yield at the expense of the nutrients in forest land. Although this genotypes grow fast, the consumption of land fertility is very high. Thus control effect on land capability decline of plantation was not obvious up till now, and it has not yet been developed effective measures to control productivity decline of continuous plantation for the production of large-scale application. How to solve the reforestation of large-scale plantation of degraded soil has now become the major issue urgent to resolve in forestry production.
This paper is based on the characteristics with low available phosphorus but rich in total phosphorus in southern forest soil, and set about improving the efficiency of the use of phosphorus of forest lands in fir through comprehensive investigation and study of growth of test stands of offspring of Chinese fir clones in different areas of different age determination and nutritional characteristics in Center of China fir. The P efficient utilization of fast-growing Chinese fir clones were initially screened out by analysis of P use efficiency in different productivity of the offspring test stands of Chinese fir clones. Through indoor simulation experiments of homogeneity and heterogeneity for the phosphorus, adaptation to physiological and biochemical differences of different low-phosphorus stress in different Chinese fir clones were analyzed and determined, then Chinese fir clones of P use efficiency were selected by increasing activation of insoluble phosphate in soil and enhancing phosphate starvation resistance, and specific indexes of P efficiency in different Chinese fir clones were pointed out, thus the internal mechanism that adapted to low-phosphorus stress with efficient utilization of P in fast-growing Chinese fir clones were clarified, ecological genetic traits of tolerance to low phosphorus nutrition of Chinese fir were revealed, so all of which provided a theoretical basis to improve the theory and methods of forest nutritional genetics; Germplasm resources of tolerance to low phosphorus of Chinese fir were developed for the promotion of production applications, which provided a new way to control the decline in continuous productivity of Chinese fir plantation. Its application prospects are extensive, and economic, ecological and social benefits are extremely significant. Main results in this paper:
1、There were significant differences in growth of progeny test plantations of the 84 different Chinese fir clones with 7a year-old. The average annual growth of tree heights、diameters at breast height and volumes of the progeny test plantations of different Chinese fir clones were 0.84 m·a~(-1)、1.12 cm·a~(-1)、0.0036 m3·a~(-1) respectively. M8, M10, M11, M23, M28, M29, M30, M32, M33, M34, M101, M102, M136, M151, M211, M239 and M249, etc 17 fast-growing Chinese fir clones were initially screened out, which accounted for 19.77 % of the total number of the tested clones.
2、There were significant differences in growth of progeny test plantations of the 45 different Chinese fir clones with 17a year-old. The average annual growth of tree heights、diameters at breast height and volumes of the progeny test plantations of different Chinese fir clones were0.94 m·a~(-1)、1.22 cm·a~(-1)、0.0161 m3·a~(-1) respectively. M17, M34, M35, M40 and M43 clones were initially screened out, which accounted for 11.11 % of the total number of the tested clones. Compared with the control 1, the average realistic genetic gains of tree heights, diameters at breast height and individual volumes in 17 clones mentioned above were 11.81 %、15.50 % and 37.16 %.
3、The average of PUE of whole tree of different Chinese fir clones with 17a year-old was 4886.67 g·g-1. Among them, PUE of whole tree was highest in M1 Chinese fir clones, which was 1.27 times of the average. Then these efficiencies were in ascending order as follows: M17>M10>M9>M45>M23>M41>M3>M19>M24>M43>M8>M25>M37>M34.PUE of stem in different clones for sequencing were: M1>M10>M9>M17>M41>M23>M19>M45>M3>M43>M8>M25>M24>M37>M34, among them, M1 was 25.84 % more than the average, which was 1.57 times of M34.
4、Different treatments of P had remarked difference on roots growth of Chinese fir clones. In rich insoluble Fe-P treatment of P1, the averages of root length, surface areas, average diameters and volumes in different Chinese fir clones were 15.58 m, 566.56 cm2, 0.77 mm and 17.13 cm3 respectively. Among them, roots in M8, M9, M11, M39 and M302 grew well, which showed that foraging ability in this 7 Chinese fir clones was stronger in the adversity of absence of available phosphorus but rich in insoluble iron and phosphorus. In P-deficiency treatment of P0, roots in M1, M3, M9, M17, M21and M302 grew well, which noted that the four Chinese fir clones had a ability of P-deficient resistance.
5、Different P treatments had remarked difference on biomass increment of Chinese fir clones. Compared with the normal phosphorus supply treatment P2, the biomass of different Chinese fir clones by 24.06 % on average in sufficient insoluble phosphate conditions in treatment P1 .Among them, increase amplitude of M302, M9, M8, M1, M11, M17, M39 and M19 were, respectively, 7.65 times, 6.23 times, 5.93 times, 4.25 times, 1.98 times, 1.84 times, 1.45 times, and 1.25 times of the average of increased percentage. It was obvious that foraging ability in this eight Chinese fir clones was stronger in the absence of available phosphorus but insoluble iron and phosphorus sufficient adversity; In P-deficiency treatment P0, biomass in different Chinese fir clones decreased by 31.97 % on average. Among them, P-deficient resistance ability was larger in M39, M1 and M302, maintaining a high productivity, and the increase in the percentage were respectively 96.17 %, 32.60 % and 26.36 % compared with that in treatment P2, which may be related to the higher phosphorus use efficiency in their body; Productivity of other tested clones was significantly influenced by P-deficient stress, and added biomass increments were smaller than that in treatment P2.
6、Different treatments of P had different impact on accumulation and distribution of P in different Chinese fir clones. The average of P content in different organs of tested 21 different Chinese fir clones showed in the order of size: leaves> roots> stems. Content of P in roots was impacted significant on low-phosphorus stress, which might be caused by those: when Chinese fir clones encountered low-phosphorus stress, On the one hand, P in plants transferred to underground portion to improve the productivity of roots to enhance the foraging ability of P in soil, on the other hand, roots activated insoluble phosphate in soil so that P content of the roots increased. In treatment of P1, P content in M5, M17, M19, M21, M24, M37, M41, M48 and M302 Chinese fir clones was higher; In treatment of P0, P Content in roots of M3, M4, M5, M19, M21, M24, M28, M39, M102 and M302 Chinese fir clones was higher.
7、Different treatments of P had different impact on P use efficiency of Chinese fir clones. In treatments of P2, P1 and P0, average of PUE in tested 21 different Chinese fir clones were 1.826 kg·g~(-1), 1.663 kg·g~(-1) and 1.814 kg·g~(-1) respectively, which further indicated that Chinese fir clones might decrease PUE mainly through activated absorption of the roots of insoluble iron phosphorus, and speeding up P recycling and reusing at the same time in the adversity of lack of available phosphorus but rich in total phosphorus; In treatment of P1, PUE in M3, M4, M19 and M38 Chinese fir clones was larger than that in treatment of P2, and the increase amplitude was 2.48 % ~ 28.04 %; In treatment of P0, PUE in M41, M1, M19, M9, M17, M11, M37, M24, M21 and M211 Chinese fir clones was all larger than that in treatment of P2, and the increase percentage was declined by 1.49 % ~ 34.94 %.
8、With the low-phosphorus stress increasing, Chlorophyll in leaves of Chinese fir decreased sooner, which showed Chlorophyll was more seriously damaged. In normal phosphorus supplied treatment of P2, as test time prolonged, average of total chlorophyll in different Chinese fir clones showed the trend that it rapidly increased firstly and then slowly increased; It showed the same law change in the treatment of P1. In treatment of P0, the total amount of chlorophyll increased rapidly at first, then rate of increase slowed down later, and decreased finally. In the end of the treatment of P1, the increase amplitude of total chlorophyll of M3, M18, M24 and M102 Chinese fir clones was larger than that in treatment of P2. In the end stress treatment of P0, only the increase amplitude of total chlorophyll of M5, M18 and M41 Chinese fir clones chlorophyll was larger than that in treatment of P2.
9、The more serious low-phosphorus stress was, the faster increased MDA content in leaves of Chinese fir. With the extension of test time, in the normal phosphorus supplied treatment of P2 ,the average MDA content in leaves of different Chinese fir clones showed such trend :it was declined slowly firstly, and then decreased substantially, finally it was slowly increased; Change law in treatment of P1 was similar to that in treatment of P2; It showed different change law in treatment of P0: there was a slow rise firstly, and then a sharp decline, finally a slow increase. Compared with the end of treatment of P2, increase amplitude of MDA content in leaves of M4, M19, M39, M102, M211 and M302 Chinese fir clones was smaller in treatment of P1. While increase amplitude of MDA content in leaves of M4, M5, M11, M18, M37, M38, M48 and M211 was smaller in treatment of P0.
10、With the stretch of test time, soluble protein content in leaves of Chinese fir clones showed similar law changes in different treatments of P: increase→decrease→slow down to the level of stress before. However, the change law of soluble protein content in leaves of Chinese fir clones was widely different. Compared with the increase amplitude of soluble protein content in leaves of Chinese fir clones in the normal phosphorus supplied treatment of P2, the increase amplitudes of M1, M3, M4, M8, M18, M19, M24, M28, M37, M38, M48 and M211 were larger in the primary stress treatment of P1, which showed that these 12 clones were more sensitive to the low-phosphorus stress. This might be due to that they could start the biological synthesis of enzymes and active substances of stress-tolerant protein in plants rapidly. At the end of stress, the increase amplitudes of M11, M18, M19, M37, M38 and M211 Chinese fir clones was larger, which might because that ability to activate insoluble iron in these six Chinese fir was larger in stress treatment of P1.This effectively alleviated the blocked plight of the normal protein synthesis in plants and improved the ability to synthesis of stress-tolerant protein. In the P-deficiency treatment of P0 at the primary stress stage, the increase amplitude of M1, M3, M4, M8, M9, M11, M19, M21, M24, M28, M38, M39, M41, M48 and M211 was larger. At the end of stress, the increase amplitude of M3, M4, M11, M18, M19, M21, M24, M38, M39 and M211 were larger.
11、With the stretch of test time, law change of the average of APA activity in leaves of Chinese fir clones in treatment of P1 and P0 was similar to the treatment of P2: decrease slowly→increase→dropped to a lower level. However, the change law of APA activity in leaves of different phosphorus efficiency of Chinese fir clones was significantly different. The primary stress stage in treatment of P1, increase amplitude of APA activity in leaves of M1, M8, M9, M11, M17, M19, M39, M48 and M211 Chinese fir clones in tested 21 Chinese fir clones was larger than that in treatment of P2; In the end of stress, the increase amplitude of M3, M5, M8, M9, M11, M18, M19, M24, M28, M37, M38, M39 and M211 Chinese fir clones was larger than that in treatment of P2.In primary low-phosphorus stress of treatment of P0, increase amplitude in APA activity in leaves of M1, M8, M38, M41 and M211 Chinese fir clones than that in normal P supplied treatment of P2; At the end of stress, increase amplitude of M1, M3, M5, M8, M18, M19, M24, M28, M37, M39, M102, M211 and M302 Chinese fir clones was larger than that in treatment of P2.
12、In different treatments of P, correlation between dry matter accumulation of Chinese fir clones and different physiological indicators was not very strong, but different morphological indicators of Chinese fir clones had more visible impact on dry matter yield. In normal phosphorus supplied treatment of P2, the correlation coefficient of the increment of root length, root surface area, root diameter and root volume of Chinese fir clones and the content of dry matter accumulation were 0.451, 0.544, 0.526 and 0.561, respectively; while the correlation coefficient were respectively 0.640, 0.686, 0.682 and 0.658 in adequate iron phosphate treatment of P1, P value test all showed significant correlation; Under phosphorus deficiency treatment of P0, root length, root area and root volume had significant impact on the dry matter yield, seedling height was also one of the decisive factor related to dry matter accumulation.
13、The study in this paper found: M39 Chinese fir clones showed strong adaptability and maintained a relatively high rate of biomass accumulation under the environment that la对照of available phosphorus but rich in total P. Use efficiency of M39 Chinese fir clones that belonged to fast-growing Chinese fir genotypes of efficient use of P was also higher; While PUE of M3, M4, M18, M19 and M38 were higher but their biomass production were lower, M1, M8, M9, M11, M17 and M302 had fast-growing and high-yield features, but relatively low PUE. It was noted that the 11 Chinese fir clones had great potential of development, which was proposed to strengthen the improvement of research.
From P-deficiency experiment of P0, this paper drew: characteristic of phosphate starvation resistance was obvious in M1 Chinese fir clones. In extreme adversity of P-deficiency, M1 Chinese fir clones could maintain a higher rate of root growth and dry matter accumulation, so it could be used as preferred material for genetic research of nutrition of Chinese fir; PUE in M9 Chinese fir clones was higher in completely P-deficiency adversity, but there was faster rate of root growth and a smaller biological accumulation. This may be related to that expanding foraging ability of the roots was higher, but the ability to reuse and transfer P in plants was poor.
14、Different heterogeneity of phosphorus supplied treatment had obvious differences of impacts on root growth of M1 and M4 Chinese fir clones. There is little difference in roots growth between 2 pots of M1 and M4 Chinese fir clones were not large. In treatments of P2, P1 and P0, increments in these four indicators of root length, surface area, mean diameter and volume in rootⅠof M1 and M4 Chinese fir clones were all larger than those in root. But with the reduction of total phosphorus in growth medium, increments of four roots growth indicators of M1 Chinese fir clones showed some increase in the trend, which might be that it could adapt to low phosphorus stress through reasonable distribution of phosphorus in plants; While the roots of M4 Chinese fir clones in the growth medium containing insoluble Fe-P grew more slowly than those in lack of total P medium, which might be related to its activation ability of the Fe-P, if phosphorus were sufficient in plants, the ability of foraging through the extension of roots weakened.
15、Heterogeneity of different P treatments had different impact on distribution of P in M1 and M4 Chinese fir clones. Differences of P content between root and stem in M1 Chinese fir clones under the normal phosphorus supplied treatment of P3 were smallest, while it had most obvious difference in P-deficiency treatment of P0; As low-phosphorus stress increased, differences in P content in roots and leaves had a descending trend. Compared with P content between rootsⅠand rootsⅡof M1Chinese fir clones, in two pots of normal phosphorus supplied treatment of P3, the difference of P content between rootⅠand rootⅡwas 0.07g·kg-1; In treatments of P2, P1 and P0, P content in rootsⅠwere respectively 0.18g·kg-1, 0.11g·kg-1 and 0.35g·kg-1 larger than those in rootsⅡ, in which two pots of treatment of P0 had the largest differences in root P content. It noted that phosphate starvation-resistant M1 Chinese fir clones could maintain normal root growth through changes in distribution pattern of phosphorus in roots Chinese fir in adversity of extremely short of phosphorus in soil.
The average of P content in roots of M4 Chinese fir clones were, respectively, 0.36 g·kg-1, 0.12 g·kg-1, 0.48 g·kg-1 and 0.10 g·kg-1 smaller than those in leaves in treatments of P3, P2, P1 and P0. Compared with P content between roots and leaves, in different treatments of P, differences were smallest in treatment of P0, while difference between the two was larger in treatment of P1. Compared with P content between rootsⅠand rootsⅡof M4 Chinese fir clones, difference of P content between rootsⅠand rootsⅡwas small by only 0.04 g·kg-1 in treatment of P3; In treatments of P2, P1 and P0, P Content in rootsⅠwere 0.11g·kg-1、0.01g·kg-1 and 0.04g·kg-1 larger than those in rootsⅡrespectively, which showed that difference of P content in roots in two pot was largest in treatment of P2.And it showed that it had a strong activation capacity of iron-phosphorus and greater phosphorus content in roots of Chinese fir clones, in the adversity of lack of available phosphorus but rich in total phosphorus in soil.
16、Heterogeneity of different P supplied treatments had obvious impact on PUE of M1 and M4 Chinese fir clones. With the increasing stress of Fe-P, PUE of M1 Chinese fir clones had a gradually increasing trend. Compared to normal phosphorus supplied treatment of P3, PUE reduced by 21.20 % and 2.94 % respectively in treatments of P2 and P1, while increased 7.78 % in treatment of P0, which noted that the less the soil P content, the higher was the efficiency of nutrient recycling in M1 Chinese fir clones and the stronger was the resistance phosphate starvation; PUE of M4 Chinese fir clones showed a slowly increasing trend, PUE reached 2.41 g·kg-1 in treatment of P0, respectively, were 12.88 %, 4.64 % and 6.54 % larger than those in treatments of P3, P1 and P0. Through comparison of PUE of M1 and M4 Chinese fir clones in different treatments of P, PUE of M4 Chinese fir clones was all larger than that of M1 Chinese fir clones, and this might be related to their different ability to activate Fe-P.
17、Different heterogeneity of phosphorus supplied treatments had obvious impact on differences of physiological indicators of M1 and M4 Chinese fir clones. The differences of physiological indicators in M1 Chinese fir clones impacted by heterogeneity of different treatments of P showed as follows: In treatment of P3, total amount of chlorophyll showed a change law of increase→slowly decrease→increase; In treatments of P3 and P0, it showed the trend of slowly increase→slowly decrease. MDA content in leaves showed a change law as follows: In treatments of P3 and P0, it showed a change trend of increase→decrease; MDA content increased slowly in treatment of P2, while MDA content showed a change trend of increase→decrease→slowly increase in treatment of P1. Soluble protein content in leaves showed a change trend that: With the prolongation of stress time, the content in all treatments rapidly increased first, and then gradually decreased. APA activity in leaves showed a change law that: in different treatments of P, it all increased rapidly after slow changes, and then weakened.
The differences of physiological indicators in M4 Chinese fir clones impacted by heterogeneity of different treatments of P showed as follows: Chlorophyll content all showed a change law in treatments of P as follows: increase→decrease→slowly increase; Among them, change range in treatment of P0 was small. MDA content in leaves showed a change law as follows: MDA content in different treatments all showed a slow increase→decrease in trend, in which decreasing range in treatment of P3 was largest and smallest in treatment of P0; Differences of change law of soluble protein content in leaves: With the prolongation of stress time, soluble protein content in different treatments all showed a change trend: rapid increased first, and then gradually decreased. Differences of change law of APA activity in leaves: it all increased rapidly after slow changes, and then weakened.
18、Heterogeneity of different P treatments had obvious impact on difference of MDA content in leaves of M1 and M4 Chinese fir clones. With the prolongation of test time, M1 Chinese fir clones showed a trend of increase→decrease in treatments of P2 and Fe0; MDA content increased slowly in treatment of Fe2, while MDA content showed a change trend of increase→decrease→slowly increase in treatment of Fe1. Before the 40 days stress, MDA content of M4 Chinese fir clones in different treatments showed a slow increase→decrease in trend, in which decreasing range in treatment of P2 was largest and smallest in treatment of Fe0; Later MDA content increased slowly in treatment of P2 and showed a decrease trend in other treatments, then MDA content in different treatments tended to the same level to the 60 days stress.
19、Different treatments of heterogeneous P had no obvious impact on difference of soluble protein content in leaves of M1 and M4 Chinese fir clones. With the prolongation of stress time, soluble protein content in different treatments all showed a change trend: rapid increased first, and then gradually decreased. Stress to 60 days, soluble protein content in leaves of M1 Chinese fir clones was largest with the value of 2.403 mg·g-1 in treatment of Fe2, which were 0.471 mg·g-1, 0.690 mg·g~(-1) and 0.365 mg·g~(-1), respectively, larger than those in treatments of P2, Fe1 and Fe0; It showed that soluble protein content in leaves of M1 and M4 Chinese fir clones was differently impacted by the low-phosphorus stress.
20、Different treatments of heterogeneous P had a little impact on differences of APA activity in leaves of M1 and M4 Chinese fir clones, all showed a change law that :increased rapidly after slow changes, and then weakened. Before the stress of 20 days, differences between different treatments was not obvious; Stress to 40 days, APA activity in M1 Chinese fir clones in treatment of P2 was strongest, up to 67.155 g~(-1)·min-1, which were 111.94 %, 55.19 % and 103.40 % larger than those in treatments of Fe2, Fe1 and Fe0.While APA activity in M4 Chinese fir clones was strongest in treatment of Fe0, which were 44.70 %, 4.55 % and 34.67 % larger than those in treatments of P2, Fe2 and Fe1; To the end of the stress, APA activity in different treatments tended to the same level.
21、In low-phosphorus stress conditions, with the prolongation of stress time, pH of hydroponics nutrient solution of Chinese fir showed a change law of rise first then fall. Compared to non-phosphorus and non-calcium treatment ofⅥ, treatment of calcium supplied helped to regulate the physiological activity of the roots of Chinese fir and promote to secrete acid materials of root. At the fourth week of low-phosphorus stress, pH value of 0.074 mol·L-1 in the treatment ofⅢ(3.0 mmol·L-1) ranked the highest level, pH value came next in treatments ofⅠ(7.0 mmol·L-1) andⅡ(5.0 mmol·L-1), respectively, 0.007 mol·L-1 and 0.008 mol·L-1 less than the former, so the differences among the three were very little; pH value in treatment ofⅥwas only 35.14 % of that in treatment ofⅢ. From the above it was clear that treatment of high-calcium supplied helped roots of Chinese fir to secrete acid materials in adversity of low-phosphorus conditions.
22、Allocation and utilization of the amount of P accumulation in Chinese fir were affected by different Ca gradient in low-phosphorus stress as follows: With the decreases of Ca gradient, the percentage accounted by the amount of P accumulation in roots reduced in order at first and then increased; the change law of leaves and stems was not obvious, but they always declined in the opposite direction. In different Ca gradient, the PUE in treatment ofⅠwas highest, amounting to 0.987 kg·g~(-1), differences in the remaining six treatments were not significant, of which PUE in CK was lowest with the value of 0.717 kg·g~(-1), it showed that the high calcium treatment had a certain impact on PUE in Chinese fir in low-phosphorus stress, but the change law was not obvious, this might be related to sustainable P-deficiency in hydroponics environment and short period of test, but this subjected to a further study.
23、The study of response of low-phosphorus stress of Chinese fir impacted by different Ca gradients showed that: In low-phosphorus stress conditions, with the prolongation of stress time, pH of hydroponics nutrient solution of Chinese fir showed a change law of rise first then fall. Compared to non-phosphorus and non-calcium treatment ofⅥ, treatment of calcium supplied helped to regulate the physiological activity of the roots of Chinese fir and promote to secrete acid materials.With the decreases of Ca gradient, the percentage accounted by the amount of P accumulation in roots reduced in order at first and then increased; the change law of leaves and stems was not obvious, but they always declined in the opposite direction. In different Ca gradient, the PUE in treatment ofⅠwas highest, amounting to 0.987 kg·g~(-1), differences in the remaining six treatments were not significant, of which PUE in CK was lowest with the value of 0.717 kg·g~(-1), it showed that the high calcium treatment had a certain impact on PUE in Chinese fir in low-phosphorus stress, but the change law was not obvious, this might be related to sustainable P-deficiency in hydroponics environment and short period of test. With the reduce of Ca concentration in hydroponics, MDA content in leaves of Chinese fir in low-phosphorus stress showed the trend as follows: decrease→increase→decrease, and the change law of total chlorophyll was similar to MDA. Change law of soluble protein content in leaves of Chinese fir was similar to that of the APA activity, the overall trend was: decrease→increase→decrease. The impact of interaction of Ca and P on response of low-phosphorus stress of Chinese fir needed a further study.
本文针对南方林区土壤有效磷低但全磷丰富的特点,从提高杉木对林地磷素的利用效率入手,通过对我国杉木中心产区不同杉木无性系子代测定林生长和磷素利用特性的全面调查,进行不同杉木无性系子代测定林生长和P素利用效率的比较研究,在此基础上初步筛选出P高效利用杉木速生无性系。利用初选的12个P高效利用速生杉木无性系为研究对象,通过均一性磷胁迫模拟试验,分析不同杉木无性系对环境低磷胁迫的形态学和生理学响应差异,探讨P高效利用杉木无性系适应低磷胁迫内在机制,进一步筛选对土壤难溶性磷活化能力高和耐低磷能强的P高效利用杉木无性系。最后利用对土壤难溶性磷活化能力高和耐低磷能强的2个P高效利用杉木无性系进行异质供磷胁迫模拟试验,通过设计不同钙梯度低磷水培处理,通过异质供磷胁迫条件下杉木无性系形态学和生理学特异性指标的测定,探讨钙在P高效利用杉木无性系对环境低磷胁迫信号感受中的作用,分析P高效利用杉木无性系适应环境磷胁迫的内在机制,为揭示杉木耐低磷营养性状的生态遗传和完善林木营养遗传学方法提供科学依据。主要的研究结果如下:
1、7年生84个不同杉木无性系子代测定林的生长存在明显差异。不同无性系子代测定林树高、胸径和材积的年均生长量分别为0.84 m·a-1、1.12 cm·a-1、0.0036 m3·a-1。初步筛选出M8、M10、M11、M23、M28、M29、M30、M32、M33、M34、M101、M102、M136、M151、M211、M239和M249等17个速生杉木无性系,占参试无性系总数的19.77 %。
2、17年生45个不同杉木无性系子代测定林的生长存在明显差异。不同杉木无性系子代测定林树高、胸径和材积的年均生长量分别为0.94 m·a-1、1.22 cm·a-1、0.0161 m3·a-1。初步筛选出了M17、M34、M35、M40和M43号等5个速生无性系,占参试无性系总数的11.11 %。与初级嫁接种子园实生苗相比,17个无性系树高、胸径和材积平均现实增益值分别为11.81 %、15.50 %和37.16 %。
3、17年生不同杉木无性系的全树P利用效率平均为4886.67 g·g-1。其中M1利用效率最高,是不同无性系平均值的1.27倍,依次排序为M17>M10>M9>M45>M23>M41>M3>M19>M24>M43>M8>M25>M37>M34。不同无性系干材P利用效率大小排序为M1>M10>M9>M17>M41>M23>M19>M45>M3>M43>M8>M25>M24>M37>M34,其中M1比平均值增加25.84 %,是M34的1.57倍。
4、初选21个P高效利用杉木速生无性系在不同P处理条件下根系生长差异明显。难溶性Fe-P充足处理中,不同杉木无性系根系长度、表面积、平均直径和体积的平均值分别为15.58 m、566.56 cm2、0.77 mm和17.13 cm3,其中M8、M9、M11、M39和M302的根系发育较好,说明这7个杉木无性系在土壤有效态磷匮乏但难溶性磷丰富的逆境中根系觅养能力较强。缺磷处理中M1、M3、M9、M17、M21和M302根系发育较好,可见这4个杉木无性系耐磷饥饿能力较强。
5、初选21个P高效利用杉木速生无性系在不同P处理条件下生物量差异较大。与正常供磷相比,难溶性磷充足处理条件下生物量平均增加24.06 %,其中M302、M9、M8、M1、M11、M17、M39和M19增幅较大,说明它们在有效磷缺乏但难溶性铁磷充足逆境中觅养能力较强,在体内抗逆机制调节作用下生物量反而高于正常供磷。缺磷处理中,不同杉木无性系生物量平均减小31.97 %,其中M39、M1和M302耐磷饥饿能力较强,保持较高生产力,这可能与其体内磷素利用效率较高有关。
6、不同P处理对初选21个P高效利用杉木无性系体内P素积累与分布影响不同。参试21个不同杉木无性系各器官P素含量平均值的高低均表现为:叶>根>茎。其中根P素含量受低磷胁迫影响显著,这可能是杉木无性系遭遇低磷胁迫时,一方面体内P素向地下部转移,提高根系生产力,以增强对土壤P素的觅养能力;另一方面是由于根系活化了土壤中难溶性磷而使根的P素含量增多。铁磷处理中,M5、M17、M19、M21、M24、M37、M41、M48和M302号杉木无性系根P素含量较高;缺磷处理中,M3、M4、M5、M19、M21、M24、M28、M39、M102和M302根P素含量较高。
7、不同P处理对不同杉木无性系P素利用效率的影响存在差异。在正常供磷、铁磷处理和缺磷处理中,参试21个杉木无性系PUE的平均值分别为1.826 kg·g-1、1.663 kg·g-1和1.814 kg·g-1。铁磷处理条件下M3、M4、M19和M38号杉木无性系PUE比正常供磷处理大,增幅为2.48 %～28.04 %;缺磷处理条件下M41、M1、M19、M9、M17、M11、M37、M24、M21和M211均比正常供磷处理大。
8、随着低磷胁迫时间的延长,参试21个不同杉木无性系叶绿素总量的平均值表现出先快速增加,再缓慢增加的趋势;铁磷处理表现出同样变化规律。缺磷处理中,叶绿素总量先迅速增加,后增加速度变缓,最后减小。铁磷处理的胁迫末期,M3、M18、M24和M102号杉木无性系叶绿素总量的增幅比正常供磷处理大。缺磷处理的胁迫末期仅有M5、M18和M41等3个杉木无性系叶绿素总量增幅比正常供磷处理大。
9、低磷胁迫越严重,杉木叶片MDA含量升高越快。随着试验时间的延长,正常供磷处理中参试21个不同杉木无性系叶片MDA含量的平均值表现为先缓慢下降,再较大幅度下降,最后缓慢增加的变化趋势;铁磷处理变化规律与正常供磷处理相似;缺磷处理表现出不同变化规律:先缓慢上升,再急剧下降,最后缓慢增加。与正常供磷处理的胁迫末期相比,铁磷处理中M4、M19、M39、M102、M211和M302叶片MDA含量增幅较小,;缺磷处理中是M4、M5、M11、M18、M37、M38、M48和M211叶片MDA含量增幅较小。
10、随着试验时间的延长,不同P处理中参试21个杉木无性系叶片可溶性蛋白含量的变化表现出相似的规律:升高→下降→缓慢下降至胁迫前水平。但不同杉木无性系叶片可溶性蛋白含量的变化规律差异悬殊。与正常供磷处理中杉木无性系叶片可溶性蛋白含量的增幅相比,铁磷处理胁迫初期,M1、M3、M4、M8、M18、M19、M24、M28、M37、M38、M48和M211号的增幅较大,表明这12个无性系对低磷胁迫较敏感,可能是由于它们能够迅速启动体内酶等抗逆蛋白活性物质的生物合成;至胁迫末期,M11、M18、M19、M37、M38和M211的增幅较大,可能是铁磷处理逆境条件下这6个杉木活化难溶性铁磷能力较强,有效缓解了体内正常蛋白质合成受阻困境,且抗逆蛋白合成能力较高。缺磷处理低磷胁迫初期时,M1、M3、M4、M8、M9、M11、M19、M21、M24、M28、M38、M39、M41、M48和M211号的增幅较大;至胁迫末期,M3、M4、M11、M18、M19、M21、M24、M38、M39和M211号的增幅较大。
11、随着试验时间的延长,铁磷处理和缺磷处理条件下初选21个P高效利用杉木速生无性系叶片APA活性的平均值的变化规律与正常供磷处理相似:缓慢下降→升高→下降至较低水平的变化规律。然而,不同杉木无性系叶片APA活性的变化规律差异显著。铁磷处理胁迫初期, M1、M8、M9、M11、M17、M19、M39、M48和M211叶片APA活性增幅比正常供磷处理大;至胁迫末期,M3、M5、M8、M9、M11、M18、M19、M24、M28、M37、M38、M39和M211比正常供磷处理大。缺磷处理低磷胁迫初期,M1、M8、M38、M41和M211叶片APA活性增幅比正常供磷处理大;胁迫末期M1、M3、M5、M8、M18、M19、M24、M28、M37、M39、M102、M211和M302比正常供磷处理大。
12、初选21个P高效利用杉木速生无性系在不同P处理条件下干物质积累与各生理学指标之间相关性不强,但与不同形态学指标密切相关。铁磷充足处理中,根长、根表面积、根直径和根体积的增量与杉木无性系干物质积累量的相关系数分别达0.640、0.686、0.682和0.658,P值检验均呈极显著相关;缺磷处理条件下,根长、根面积和根体积对干物质产量影响显著,苗高也是决定干物质积累的相关因子之一。PUE与不同形态学指标相关性不大,不同生理学指标中叶绿素a和叶绿素总量均与杉木无性系PUE呈显著负相关。铁磷处理中,杉无性系PUE与叶片可溶性蛋白含量和APA活性分别呈显著水平和极显著水平,说明在林地有效磷匮乏但全磷丰富条件下,叶片的可溶性蛋白含量及APA活性可以作为评价筛选P素高效利用杉木无性系的可靠生理指标。
13、参试21个杉木无性系中M39号杉木无性系在有效磷匮乏全磷丰富的环境中适应性较强,且能保持较高的生物量积累速度,对P素的利用效率也较高,属于P高效利用杉木速生基因型;而M3、M4、M18、M19和M38的PUE较高但生物量产量较低,M1、M8、M9、M11、M17和M302具速生丰产特性,但PUE较低,可见这11个杉木无性系的开发潜力大,建议加强改良研究。缺磷处理试验表明:M1号杉木无性系耐磷饥饿特性明显,在极度缺磷的逆境中,能维持较高的根生长和干物质积累速度,可作为杉木营养遗传学研究的首选材料;M9号杉木无性系在完全缺P逆境中PUE较高,根生长速度较快,但生物积累量较小,这可能是与其通过扩大根系觅养能力较强,但体内P素的转移重复利用的能力较差有关。
14、不同异质供磷处理对M1和M4号杉木无性系根系生长的影响差异明显。两室均为正常磷供处理中,M1和M4两室根系生长差异均不大。高浓度铁磷、低浓度铁磷和缺磷处理条件下,M1和M4室Ⅰ中的根系长度、表面积、平均直径和体积生长量均比室Ⅱ大,但随着生长介质中磷素全量的减少,M1根系长度、表面积、平均直径和体积生长量均呈一定增加的趋势;而M4在生长介质中含有难溶性铁磷时根系生长速度较完全缺P时慢,这可能与它活化铁磷能力较强有关,一旦体内磷素充足,其通过根系延伸进行觅养的能力减弱。
15、不同异质P处理对M1和M4号杉木无性系体内P素分布的影响不同。M1号杉木无性系正常供磷处理条件下根与茎的P素含量相差最小,而在缺磷处理中差异最为明显;随着低磷胁迫加重,根与叶P素含量的差值递减趋势。通过对M1号杉木无性系室Ⅰ和室Ⅱ中根P素含量的比较,高浓度铁磷、低浓度铁磷和缺磷处理条件下室Ⅰ的根P素含量分别比室Ⅱ大0.18g·kg~(-1)、0.11g·kg~(-1)和0.35g·kg~(-1),表明M1号杉木在土壤磷极度匮乏逆境中耐磷饥饿杉木无性系可通过根系磷素分配格局的变化来维持根系的正常生长。
M4号杉木无性系正常供磷、高浓度铁磷、低浓度铁磷和缺磷处理根系平均P素含量分别比叶小了0.36 g·kg~(-1)、0.12 g·kg~(-1)、0.48 g·kg~(-1)和0.10 g·kg~(-1)。根与叶P素含量相比,不同P处理中缺磷处理中两者差异最小,而低浓度铁磷处理中两者差异较大。通过对M4号杉木无性系室Ⅰ和室Ⅱ中根P素含量的比较,高浓度铁磷、低浓度铁磷和缺磷处理条件下室Ⅰ中根的P素含量分别比室Ⅱ大0.11 g·kg~(-1)、0.01 g·kg~(-1)和0.04 g·kg~(-1),表明高浓度铁磷处理中两室根系P含量差异最大,说明在土壤有效磷匮乏但全磷丰富逆境中,活化铁磷能力较强杉木无性系根系磷素含量较大。
16、不同异质供磷处理对M1号和M4号杉木无性系PUE的影响差异较大。随着铁磷胁迫加重,M1号杉木无性系PUE呈逐渐升高趋势,与正常供磷处理相比,高浓度铁磷处理和低浓度铁磷处理PUE的值分别减小了21.20 %和2.94 %,而P0处理增大了7.78 %,说明土壤P含量越少,M1体内养分回收利用效率越高,耐磷饥饿越强;M4号杉木无性系PUE呈缓慢升高趋势,缺磷处理中PUE达2.41 kg·g~(-1),分别比正常供磷处理、高浓度铁磷和低浓度铁磷处理大了12.88 %、6.54 %和4.64 %。通过不同P处理M1和M4号杉木无性系PUE的比较,M4号杉木无性系PUE均比M1大,这可能与它们对铁磷活化能力不同有关。
17、不同异质供P处理对M1和M4号杉木无性系各生理指标的影响差异明显。不同异质供P处理对M1号杉木无性系各生理指标影响的差异表现为:正常供磷处理叶绿素总量表现出升高→缓慢降低→升高的变化规律;高浓度铁磷处理和缺磷处理呈缓慢升高→缓慢降低的变化趋势。叶片MDA含量变化规律的差异:正常供磷和缺磷处理条件下呈升高→降低的变化趋势;高浓度铁磷处理缓慢增加,而低浓度铁磷处理表现出升高→降低→缓慢升高的变化规律。叶片可溶性蛋白含量变化规律的差异:不同处理均随着胁迫时间的延长,表现出先快速升高,再逐渐降低的变化趋势。叶片APA活性变化规律的差异:不同P处理均表现为缓慢变化之后,快速增强,再减弱的变化趋势。
不同异质供P处理对M4号杉木无性系各生理指标影响的差异表现为:不同P处理叶绿素含量均表现出升高→降低→缓慢升高的变化规律,其中缺磷处理变化幅度很小。叶片MDA含量变化规律的差异:不同处理均表现出缓慢升高→降低的变化趋势,其中正常供磷处理降幅最大,缺磷处理最小。叶片可溶性蛋白含量变化规律的差异:不同处理均随着胁迫时间的延长,表现出先快速升高,再逐渐降低的变化趋势。叶片APA活性变化规律的差异:不同P处理均表现为缓慢变化之后,快速增强,再减弱的变化趋势。
18、不同Ca梯度在杉木感受低磷胁迫的试验表明:低磷胁迫条件下随胁迫时间的延长,水培杉木营养液酸度呈先上升后下降的变化规律。与无磷无钙处理相比,加钙处理有助于调节杉木根系生理活性,促进根系分泌酸性物质。随着Ca梯度的减小,杉木根P积累量所占百分数先依次降低再升高;叶和茎的变化规律不明显,但两者总是呈相反方向增减。不同Ca梯度中,处理Ⅰ的PUE最高,其余6个处理差异未达显著水平,其中对照最低,值为0.717 kg·g~(-1),表明高钙处理对低磷胁迫杉木PUE有一定的影响,但变化规律不明显,这可能与水培环境持续缺磷且试验培养期较短有关。不同Ca梯度处理条件下,低磷胁迫杉木各生理学指标的变化规律不明显。随着水培施Ca浓度降低,低磷胁迫杉木叶片MDA含量呈先下降后升高再降低的变化趋势,叶绿素总量的变化规律与MDA相近;叶片可溶性蛋白含量与APA活性的变化规律相似,总体趋势表现为降低→升高→降低。有关Ca和P的互作在杉木感受环境低磷胁迫中的作用还需进一步深入研究。
Chinese fir (Cunninghamia Lanceolata) is the unique important fast-growing timber species in southern China. With the growing planting area of Chinese fir, broadleaved forests area reduced sharply, more and more Chinese fir planted continuously at the same ground, which caused a decline in plantation productivity, so that the arisen "second generation effect" of plantation management had a serious impact on the continued operation of Chinese fir plantation. A large area of the fast growing Chinese fir forest has built during the 1960s and 1970s in southern China forests, most of which have entered into the final cutting stage and all faced with the problem of second-generation update. Thus settlement of soil degradation caused by second- generation update of large area Chinese fir plantation in the South has become the major subject,which starve for solution in the current forestry production. At present, prevention and control measures of soil degradation in Chinese fir plantation mainly promote the growth of Chinese fir through the improvement of soil conditions.This traditional control methods of changing land with trees are with little regard for the efficiency of nutrient use by selecting the high forest trees to adapt to the soil conditions to study, especially efficient use of nutrition genotypes are rarely researched to gain high yields at the lower level of fertility in forest land. The current fast-growing genotypes of forest trees is not necessarily the highest in nutrient use efficiency, some get high yield at the expense of the nutrients in forest land. Although this genotypes grow fast, the consumption of land fertility is very high. Thus control effect on land capability decline of plantation was not obvious up till now, and it has not yet been developed effective measures to control productivity decline of continuous plantation for the production of large-scale application. How to solve the reforestation of large-scale plantation of degraded soil has now become the major issue urgent to resolve in forestry production.
This paper is based on the characteristics with low available phosphorus but rich in total phosphorus in southern forest soil, and set about improving the efficiency of the use of phosphorus of forest lands in fir through comprehensive investigation and study of growth of test stands of offspring of Chinese fir clones in different areas of different age determination and nutritional characteristics in Center of China fir. The P efficient utilization of fast-growing Chinese fir clones were initially screened out by analysis of P use efficiency in different productivity of the offspring test stands of Chinese fir clones. Through indoor simulation experiments of homogeneity and heterogeneity for the phosphorus, adaptation to physiological and biochemical differences of different low-phosphorus stress in different Chinese fir clones were analyzed and determined, then Chinese fir clones of P use efficiency were selected by increasing activation of insoluble phosphate in soil and enhancing phosphate starvation resistance, and specific indexes of P efficiency in different Chinese fir clones were pointed out, thus the internal mechanism that adapted to low-phosphorus stress with efficient utilization of P in fast-growing Chinese fir clones were clarified, ecological genetic traits of tolerance to low phosphorus nutrition of Chinese fir were revealed, so all of which provided a theoretical basis to improve the theory and methods of forest nutritional genetics; Germplasm resources of tolerance to low phosphorus of Chinese fir were developed for the promotion of production applications, which provided a new way to control the decline in continuous productivity of Chinese fir plantation. Its application prospects are extensive, and economic, ecological and social benefits are extremely significant. Main results in this paper:
1、There were significant differences in growth of progeny test plantations of the 84 different Chinese fir clones with 7a year-old. The average annual growth of tree heights、diameters at breast height and volumes of the progeny test plantations of different Chinese fir clones were 0.84 m·a~(-1)、1.12 cm·a~(-1)、0.0036 m3·a~(-1) respectively. M8, M10, M11, M23, M28, M29, M30, M32, M33, M34, M101, M102, M136, M151, M211, M239 and M249, etc 17 fast-growing Chinese fir clones were initially screened out, which accounted for 19.77 % of the total number of the tested clones.
2、There were significant differences in growth of progeny test plantations of the 45 different Chinese fir clones with 17a year-old. The average annual growth of tree heights、diameters at breast height and volumes of the progeny test plantations of different Chinese fir clones were0.94 m·a~(-1)、1.22 cm·a~(-1)、0.0161 m3·a~(-1) respectively. M17, M34, M35, M40 and M43 clones were initially screened out, which accounted for 11.11 % of the total number of the tested clones. Compared with the control 1, the average realistic genetic gains of tree heights, diameters at breast height and individual volumes in 17 clones mentioned above were 11.81 %、15.50 % and 37.16 %.
3、The average of PUE of whole tree of different Chinese fir clones with 17a year-old was 4886.67 g·g-1. Among them, PUE of whole tree was highest in M1 Chinese fir clones, which was 1.27 times of the average. Then these efficiencies were in ascending order as follows: M17>M10>M9>M45>M23>M41>M3>M19>M24>M43>M8>M25>M37>M34.PUE of stem in different clones for sequencing were: M1>M10>M9>M17>M41>M23>M19>M45>M3>M43>M8>M25>M24>M37>M34, among them, M1 was 25.84 % more than the average, which was 1.57 times of M34.
4、Different treatments of P had remarked difference on roots growth of Chinese fir clones. In rich insoluble Fe-P treatment of P1, the averages of root length, surface areas, average diameters and volumes in different Chinese fir clones were 15.58 m, 566.56 cm2, 0.77 mm and 17.13 cm3 respectively. Among them, roots in M8, M9, M11, M39 and M302 grew well, which showed that foraging ability in this 7 Chinese fir clones was stronger in the adversity of absence of available phosphorus but rich in insoluble iron and phosphorus. In P-deficiency treatment of P0, roots in M1, M3, M9, M17, M21and M302 grew well, which noted that the four Chinese fir clones had a ability of P-deficient resistance.
5、Different P treatments had remarked difference on biomass increment of Chinese fir clones. Compared with the normal phosphorus supply treatment P2, the biomass of different Chinese fir clones by 24.06 % on average in sufficient insoluble phosphate conditions in treatment P1 .Among them, increase amplitude of M302, M9, M8, M1, M11, M17, M39 and M19 were, respectively, 7.65 times, 6.23 times, 5.93 times, 4.25 times, 1.98 times, 1.84 times, 1.45 times, and 1.25 times of the average of increased percentage. It was obvious that foraging ability in this eight Chinese fir clones was stronger in the absence of available phosphorus but insoluble iron and phosphorus sufficient adversity; In P-deficiency treatment P0, biomass in different Chinese fir clones decreased by 31.97 % on average. Among them, P-deficient resistance ability was larger in M39, M1 and M302, maintaining a high productivity, and the increase in the percentage were respectively 96.17 %, 32.60 % and 26.36 % compared with that in treatment P2, which may be related to the higher phosphorus use efficiency in their body; Productivity of other tested clones was significantly influenced by P-deficient stress, and added biomass increments were smaller than that in treatment P2.
6、Different treatments of P had different impact on accumulation and distribution of P in different Chinese fir clones. The average of P content in different organs of tested 21 different Chinese fir clones showed in the order of size: leaves> roots> stems. Content of P in roots was impacted significant on low-phosphorus stress, which might be caused by those: when Chinese fir clones encountered low-phosphorus stress, On the one hand, P in plants transferred to underground portion to improve the productivity of roots to enhance the foraging ability of P in soil, on the other hand, roots activated insoluble phosphate in soil so that P content of the roots increased. In treatment of P1, P content in M5, M17, M19, M21, M24, M37, M41, M48 and M302 Chinese fir clones was higher; In treatment of P0, P Content in roots of M3, M4, M5, M19, M21, M24, M28, M39, M102 and M302 Chinese fir clones was higher.
7、Different treatments of P had different impact on P use efficiency of Chinese fir clones. In treatments of P2, P1 and P0, average of PUE in tested 21 different Chinese fir clones were 1.826 kg·g~(-1), 1.663 kg·g~(-1) and 1.814 kg·g~(-1) respectively, which further indicated that Chinese fir clones might decrease PUE mainly through activated absorption of the roots of insoluble iron phosphorus, and speeding up P recycling and reusing at the same time in the adversity of lack of available phosphorus but rich in total phosphorus; In treatment of P1, PUE in M3, M4, M19 and M38 Chinese fir clones was larger than that in treatment of P2, and the increase amplitude was 2.48 % ~ 28.04 %; In treatment of P0, PUE in M41, M1, M19, M9, M17, M11, M37, M24, M21 and M211 Chinese fir clones was all larger than that in treatment of P2, and the increase percentage was declined by 1.49 % ~ 34.94 %.
8、With the low-phosphorus stress increasing, Chlorophyll in leaves of Chinese fir decreased sooner, which showed Chlorophyll was more seriously damaged. In normal phosphorus supplied treatment of P2, as test time prolonged, average of total chlorophyll in different Chinese fir clones showed the trend that it rapidly increased firstly and then slowly increased; It showed the same law change in the treatment of P1. In treatment of P0, the total amount of chlorophyll increased rapidly at first, then rate of increase slowed down later, and decreased finally. In the end of the treatment of P1, the increase amplitude of total chlorophyll of M3, M18, M24 and M102 Chinese fir clones was larger than that in treatment of P2. In the end stress treatment of P0, only the increase amplitude of total chlorophyll of M5, M18 and M41 Chinese fir clones chlorophyll was larger than that in treatment of P2.
9、The more serious low-phosphorus stress was, the faster increased MDA content in leaves of Chinese fir. With the extension of test time, in the normal phosphorus supplied treatment of P2 ,the average MDA content in leaves of different Chinese fir clones showed such trend :it was declined slowly firstly, and then decreased substantially, finally it was slowly increased; Change law in treatment of P1 was similar to that in treatment of P2; It showed different change law in treatment of P0: there was a slow rise firstly, and then a sharp decline, finally a slow increase. Compared with the end of treatment of P2, increase amplitude of MDA content in leaves of M4, M19, M39, M102, M211 and M302 Chinese fir clones was smaller in treatment of P1. While increase amplitude of MDA content in leaves of M4, M5, M11, M18, M37, M38, M48 and M211 was smaller in treatment of P0.
10、With the stretch of test time, soluble protein content in leaves of Chinese fir clones showed similar law changes in different treatments of P: increase→decrease→slow down to the level of stress before. However, the change law of soluble protein content in leaves of Chinese fir clones was widely different. Compared with the increase amplitude of soluble protein content in leaves of Chinese fir clones in the normal phosphorus supplied treatment of P2, the increase amplitudes of M1, M3, M4, M8, M18, M19, M24, M28, M37, M38, M48 and M211 were larger in the primary stress treatment of P1, which showed that these 12 clones were more sensitive to the low-phosphorus stress. This might be due to that they could start the biological synthesis of enzymes and active substances of stress-tolerant protein in plants rapidly. At the end of stress, the increase amplitudes of M11, M18, M19, M37, M38 and M211 Chinese fir clones was larger, which might because that ability to activate insoluble iron in these six Chinese fir was larger in stress treatment of P1.This effectively alleviated the blocked plight of the normal protein synthesis in plants and improved the ability to synthesis of stress-tolerant protein. In the P-deficiency treatment of P0 at the primary stress stage, the increase amplitude of M1, M3, M4, M8, M9, M11, M19, M21, M24, M28, M38, M39, M41, M48 and M211 was larger. At the end of stress, the increase amplitude of M3, M4, M11, M18, M19, M21, M24, M38, M39 and M211 were larger.
11、With the stretch of test time, law change of the average of APA activity in leaves of Chinese fir clones in treatment of P1 and P0 was similar to the treatment of P2: decrease slowly→increase→dropped to a lower level. However, the change law of APA activity in leaves of different phosphorus efficiency of Chinese fir clones was significantly different. The primary stress stage in treatment of P1, increase amplitude of APA activity in leaves of M1, M8, M9, M11, M17, M19, M39, M48 and M211 Chinese fir clones in tested 21 Chinese fir clones was larger than that in treatment of P2; In the end of stress, the increase amplitude of M3, M5, M8, M9, M11, M18, M19, M24, M28, M37, M38, M39 and M211 Chinese fir clones was larger than that in treatment of P2.In primary low-phosphorus stress of treatment of P0, increase amplitude in APA activity in leaves of M1, M8, M38, M41 and M211 Chinese fir clones than that in normal P supplied treatment of P2; At the end of stress, increase amplitude of M1, M3, M5, M8, M18, M19, M24, M28, M37, M39, M102, M211 and M302 Chinese fir clones was larger than that in treatment of P2.
12、In different treatments of P, correlation between dry matter accumulation of Chinese fir clones and different physiological indicators was not very strong, but different morphological indicators of Chinese fir clones had more visible impact on dry matter yield. In normal phosphorus supplied treatment of P2, the correlation coefficient of the increment of root length, root surface area, root diameter and root volume of Chinese fir clones and the content of dry matter accumulation were 0.451, 0.544, 0.526 and 0.561, respectively; while the correlation coefficient were respectively 0.640, 0.686, 0.682 and 0.658 in adequate iron phosphate treatment of P1, P value test all showed significant correlation; Under phosphorus deficiency treatment of P0, root length, root area and root volume had significant impact on the dry matter yield, seedling height was also one of the decisive factor related to dry matter accumulation.
13、The study in this paper found: M39 Chinese fir clones showed strong adaptability and maintained a relatively high rate of biomass accumulation under the environment that la对照of available phosphorus but rich in total P. Use efficiency of M39 Chinese fir clones that belonged to fast-growing Chinese fir genotypes of efficient use of P was also higher; While PUE of M3, M4, M18, M19 and M38 were higher but their biomass production were lower, M1, M8, M9, M11, M17 and M302 had fast-growing and high-yield features, but relatively low PUE. It was noted that the 11 Chinese fir clones had great potential of development, which was proposed to strengthen the improvement of research.
From P-deficiency experiment of P0, this paper drew: characteristic of phosphate starvation resistance was obvious in M1 Chinese fir clones. In extreme adversity of P-deficiency, M1 Chinese fir clones could maintain a higher rate of root growth and dry matter accumulation, so it could be used as preferred material for genetic research of nutrition of Chinese fir; PUE in M9 Chinese fir clones was higher in completely P-deficiency adversity, but there was faster rate of root growth and a smaller biological accumulation. This may be related to that expanding foraging ability of the roots was higher, but the ability to reuse and transfer P in plants was poor.
14、Different heterogeneity of phosphorus supplied treatment had obvious differences of impacts on root growth of M1 and M4 Chinese fir clones. There is little difference in roots growth between 2 pots of M1 and M4 Chinese fir clones were not large. In treatments of P2, P1 and P0, increments in these four indicators of root length, surface area, mean diameter and volume in rootⅠof M1 and M4 Chinese fir clones were all larger than those in root. But with the reduction of total phosphorus in growth medium, increments of four roots growth indicators of M1 Chinese fir clones showed some increase in the trend, which might be that it could adapt to low phosphorus stress through reasonable distribution of phosphorus in plants; While the roots of M4 Chinese fir clones in the growth medium containing insoluble Fe-P grew more slowly than those in lack of total P medium, which might be related to its activation ability of the Fe-P, if phosphorus were sufficient in plants, the ability of foraging through the extension of roots weakened.
15、Heterogeneity of different P treatments had different impact on distribution of P in M1 and M4 Chinese fir clones. Differences of P content between root and stem in M1 Chinese fir clones under the normal phosphorus supplied treatment of P3 were smallest, while it had most obvious difference in P-deficiency treatment of P0; As low-phosphorus stress increased, differences in P content in roots and leaves had a descending trend. Compared with P content between rootsⅠand rootsⅡof M1Chinese fir clones, in two pots of normal phosphorus supplied treatment of P3, the difference of P content between rootⅠand rootⅡwas 0.07g·kg-1; In treatments of P2, P1 and P0, P content in rootsⅠwere respectively 0.18g·kg-1, 0.11g·kg-1 and 0.35g·kg-1 larger than those in rootsⅡ, in which two pots of treatment of P0 had the largest differences in root P content. It noted that phosphate starvation-resistant M1 Chinese fir clones could maintain normal root growth through changes in distribution pattern of phosphorus in roots Chinese fir in adversity of extremely short of phosphorus in soil.
The average of P content in roots of M4 Chinese fir clones were, respectively, 0.36 g·kg-1, 0.12 g·kg-1, 0.48 g·kg-1 and 0.10 g·kg-1 smaller than those in leaves in treatments of P3, P2, P1 and P0. Compared with P content between roots and leaves, in different treatments of P, differences were smallest in treatment of P0, while difference between the two was larger in treatment of P1. Compared with P content between rootsⅠand rootsⅡof M4 Chinese fir clones, difference of P content between rootsⅠand rootsⅡwas small by only 0.04 g·kg-1 in treatment of P3; In treatments of P2, P1 and P0, P Content in rootsⅠwere 0.11g·kg-1、0.01g·kg-1 and 0.04g·kg-1 larger than those in rootsⅡrespectively, which showed that difference of P content in roots in two pot was largest in treatment of P2.And it showed that it had a strong activation capacity of iron-phosphorus and greater phosphorus content in roots of Chinese fir clones, in the adversity of lack of available phosphorus but rich in total phosphorus in soil.
16、Heterogeneity of different P supplied treatments had obvious impact on PUE of M1 and M4 Chinese fir clones. With the increasing stress of Fe-P, PUE of M1 Chinese fir clones had a gradually increasing trend. Compared to normal phosphorus supplied treatment of P3, PUE reduced by 21.20 % and 2.94 % respectively in treatments of P2 and P1, while increased 7.78 % in treatment of P0, which noted that the less the soil P content, the higher was the efficiency of nutrient recycling in M1 Chinese fir clones and the stronger was the resistance phosphate starvation; PUE of M4 Chinese fir clones showed a slowly increasing trend, PUE reached 2.41 g·kg-1 in treatment of P0, respectively, were 12.88 %, 4.64 % and 6.54 % larger than those in treatments of P3, P1 and P0. Through comparison of PUE of M1 and M4 Chinese fir clones in different treatments of P, PUE of M4 Chinese fir clones was all larger than that of M1 Chinese fir clones, and this might be related to their different ability to activate Fe-P.
17、Different heterogeneity of phosphorus supplied treatments had obvious impact on differences of physiological indicators of M1 and M4 Chinese fir clones. The differences of physiological indicators in M1 Chinese fir clones impacted by heterogeneity of different treatments of P showed as follows: In treatment of P3, total amount of chlorophyll showed a change law of increase→slowly decrease→increase; In treatments of P3 and P0, it showed the trend of slowly increase→slowly decrease. MDA content in leaves showed a change law as follows: In treatments of P3 and P0, it showed a change trend of increase→decrease; MDA content increased slowly in treatment of P2, while MDA content showed a change trend of increase→decrease→slowly increase in treatment of P1. Soluble protein content in leaves showed a change trend that: With the prolongation of stress time, the content in all treatments rapidly increased first, and then gradually decreased. APA activity in leaves showed a change law that: in different treatments of P, it all increased rapidly after slow changes, and then weakened.
The differences of physiological indicators in M4 Chinese fir clones impacted by heterogeneity of different treatments of P showed as follows: Chlorophyll content all showed a change law in treatments of P as follows: increase→decrease→slowly increase; Among them, change range in treatment of P0 was small. MDA content in leaves showed a change law as follows: MDA content in different treatments all showed a slow increase→decrease in trend, in which decreasing range in treatment of P3 was largest and smallest in treatment of P0; Differences of change law of soluble protein content in leaves: With the prolongation of stress time, soluble protein content in different treatments all showed a change trend: rapid increased first, and then gradually decreased. Differences of change law of APA activity in leaves: it all increased rapidly after slow changes, and then weakened.
18、Heterogeneity of different P treatments had obvious impact on difference of MDA content in leaves of M1 and M4 Chinese fir clones. With the prolongation of test time, M1 Chinese fir clones showed a trend of increase→decrease in treatments of P2 and Fe0; MDA content increased slowly in treatment of Fe2, while MDA content showed a change trend of increase→decrease→slowly increase in treatment of Fe1. Before the 40 days stress, MDA content of M4 Chinese fir clones in different treatments showed a slow increase→decrease in trend, in which decreasing range in treatment of P2 was largest and smallest in treatment of Fe0; Later MDA content increased slowly in treatment of P2 and showed a decrease trend in other treatments, then MDA content in different treatments tended to the same level to the 60 days stress.
19、Different treatments of heterogeneous P had no obvious impact on difference of soluble protein content in leaves of M1 and M4 Chinese fir clones. With the prolongation of stress time, soluble protein content in different treatments all showed a change trend: rapid increased first, and then gradually decreased. Stress to 60 days, soluble protein content in leaves of M1 Chinese fir clones was largest with the value of 2.403 mg·g-1 in treatment of Fe2, which were 0.471 mg·g-1, 0.690 mg·g~(-1) and 0.365 mg·g~(-1), respectively, larger than those in treatments of P2, Fe1 and Fe0; It showed that soluble protein content in leaves of M1 and M4 Chinese fir clones was differently impacted by the low-phosphorus stress.
20、Different treatments of heterogeneous P had a little impact on differences of APA activity in leaves of M1 and M4 Chinese fir clones, all showed a change law that :increased rapidly after slow changes, and then weakened. Before the stress of 20 days, differences between different treatments was not obvious; Stress to 40 days, APA activity in M1 Chinese fir clones in treatment of P2 was strongest, up to 67.155 g~(-1)·min-1, which were 111.94 %, 55.19 % and 103.40 % larger than those in treatments of Fe2, Fe1 and Fe0.While APA activity in M4 Chinese fir clones was strongest in treatment of Fe0, which were 44.70 %, 4.55 % and 34.67 % larger than those in treatments of P2, Fe2 and Fe1; To the end of the stress, APA activity in different treatments tended to the same level.
21、In low-phosphorus stress conditions, with the prolongation of stress time, pH of hydroponics nutrient solution of Chinese fir showed a change law of rise first then fall. Compared to non-phosphorus and non-calcium treatment ofⅥ, treatment of calcium supplied helped to regulate the physiological activity of the roots of Chinese fir and promote to secrete acid materials of root. At the fourth week of low-phosphorus stress, pH value of 0.074 mol·L-1 in the treatment ofⅢ(3.0 mmol·L-1) ranked the highest level, pH value came next in treatments ofⅠ(7.0 mmol·L-1) andⅡ(5.0 mmol·L-1), respectively, 0.007 mol·L-1 and 0.008 mol·L-1 less than the former, so the differences among the three were very little; pH value in treatment ofⅥwas only 35.14 % of that in treatment ofⅢ. From the above it was clear that treatment of high-calcium supplied helped roots of Chinese fir to secrete acid materials in adversity of low-phosphorus conditions.
22、Allocation and utilization of the amount of P accumulation in Chinese fir were affected by different Ca gradient in low-phosphorus stress as follows: With the decreases of Ca gradient, the percentage accounted by the amount of P accumulation in roots reduced in order at first and then increased; the change law of leaves and stems was not obvious, but they always declined in the opposite direction. In different Ca gradient, the PUE in treatment ofⅠwas highest, amounting to 0.987 kg·g~(-1), differences in the remaining six treatments were not significant, of which PUE in CK was lowest with the value of 0.717 kg·g~(-1), it showed that the high calcium treatment had a certain impact on PUE in Chinese fir in low-phosphorus stress, but the change law was not obvious, this might be related to sustainable P-deficiency in hydroponics environment and short period of test, but this subjected to a further study.
23、The study of response of low-phosphorus stress of Chinese fir impacted by different Ca gradients showed that: In low-phosphorus stress conditions, with the prolongation of stress time, pH of hydroponics nutrient solution of Chinese fir showed a change law of rise first then fall. Compared to non-phosphorus and non-calcium treatment ofⅥ, treatment of calcium supplied helped to regulate the physiological activity of the roots of Chinese fir and promote to secrete acid materials.With the decreases of Ca gradient, the percentage accounted by the amount of P accumulation in roots reduced in order at first and then increased; the change law of leaves and stems was not obvious, but they always declined in the opposite direction. In different Ca gradient, the PUE in treatment ofⅠwas highest, amounting to 0.987 kg·g~(-1), differences in the remaining six treatments were not significant, of which PUE in CK was lowest with the value of 0.717 kg·g~(-1), it showed that the high calcium treatment had a certain impact on PUE in Chinese fir in low-phosphorus stress, but the change law was not obvious, this might be related to sustainable P-deficiency in hydroponics environment and short period of test. With the reduce of Ca concentration in hydroponics, MDA content in leaves of Chinese fir in low-phosphorus stress showed the trend as follows: decrease→increase→decrease, and the change law of total chlorophyll was similar to MDA. Change law of soluble protein content in leaves of Chinese fir was similar to that of the APA activity, the overall trend was: decrease→increase→decrease. The impact of interaction of Ca and P on response of low-phosphorus stress of Chinese fir needed a further study.
引文
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