不同发育阶段杉木人工林养分内循环与周转利用效率的研究
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摘要
近些年来杉木人工林多代连栽生产力下降日趋明显,但目前对杉木连栽地力衰退的内在机制还不很清楚,因此,如何进一步揭示杉木人工林连栽地力衰退的内在机制成为当前林业生产中急需解决的重大课题。广大研究者将杉木人工林地力衰退的主要原因归结为炼山、皆伐、除草等各种不合理营林措施,而较少从杉木自身的生物学特性及养分利用状况进行探讨。凋落物和细根是人工林生态系统的重要组成部分,是养分利用的核心,凋落物的产量及细根生长、周转速率是影响林地土壤肥力的重要因素。虽然有关杉木人工林的凋落物特性、细根周转和养分循环进行了大量研究,并取得了一定成果,但关于从凋落物及细根养分内循环角度系统地探讨杉木人工林养分利用规律的报导还比较少,未能揭示出杉木人工林地力衰退的根本原因和内在机制。
有鉴于此,本研究针对杉木人工林养分循环研究中存在的问题,从养分内循环和周转利用效率入手,在福建农林大学莘口教学林场杉木人工林定位观测站的林分内,选择不同发育阶段的杉木幼龄林、成龄林和过熟林作为研究对象,研究不同发育阶段杉木人工林凋落物数量、养分归还、养分转移以及细根生长和衰老过程的养分转移,分析了不同发育阶段杉木林养分周转和利用效率,对提高杉木人工林生态系统的养分利用率和生产力水平具有重要现实意义,为深入揭示杉木人工林地力衰退的内在机制提供理论依据。主要研究结果如下：
1、不同发育阶段杉木人工林生物量、生产力水平及养分利用效率存在一定差异。生物量与生产力水平均表现为随发育阶段的增加而增加,幼龄林、成龄林和过熟林乔木层生物量分别为38.11、104.03和138.24t·hm-2,生产力分别为4.80、6.97和8.11t·hm-2·a-1。养分利用效率表现为随发育阶段的增加而增加,具体表现为林全树生产1吨干物质需要的大量养分量分别为18.58、13.66和12.74kg,微量养分量分别为0.608、0.490和0.490kg；大量养分利用效率表现为P>Mg>Ca>K>N,微量养分利用效率表现为Cu>Zn>Fe或Mn。通过比较各养分循环参数指标,表明幼龄林杉木具有高吸收、低归还、周转时间短、养分利用率低的特点,成熟林杉木养分循环系数提高,养分周转期延长,具有高吸收、高归还、养分消耗大特点,过熟林养分周转期延长,具有低吸收、高归还、养分利用效率高的特点。杉木生长后期更多关注自身生长,通过提高养分利用效率减少对地力的压力,若在杉木幼龄林时进行收获,势必带走大量养分,影响整个生态系统的平衡,因此,适当延长杉木轮伐期,对恢复地力意义重大,
2、通过对凋落物收集方式的筛选研究,结果表明：不同布点高度对收集量的影响表现为50cm>25cm>100cm;不同布点方式表现为“随机”>“×”>“+”；不同收集器面积表现为0.5m2>2m2>1m2(p<0.05)；收集器总面积占标准地的比例表现为1.5%>0.75%>0.5%。2012-2013年均凋落总量表现为随发育阶段的增加而增加,即过熟林(507.84g·m-2·a-1)>成龄林(458.79g·m-2·a-1)>幼龄林(348.52g·m-2·a-1),但具有年季变化特征。杉木凋落物各组分之间及其与总量之间存在异速比例关系,可以通过异速生长方程估测各组分的凋落量。叶、枝、花果、其它组分与总凋落量异速指数分别为1.36、1.41、1.52和1.31；叶与枝、叶与花果、枝与花果凋落量异速指数均小于1,分别为0.91、0.82和0.84,叶与其它、枝与其它和花与其它凋落量异速指均大于1,分别为1.19、1.19和1.32。凋落物各组分所占比例不同,落叶所占比例最大(32.9%-44.4%),枝次之(15.2%-17.2%),花果所占比例最小(2.4%-10.8%)。杉木人工林凋落物产量具有明显的季节节律,总体而言,凋落量出现在4-5月、8月和12月,针叶与枝凋落量月动态与总凋落量变化节律相似;季节变化表现为春季>夏季>秋季>冬季,叶主要集中在春季凋落,枝、花果凋落主要集中在夏季凋落。
不同发育阶段杉木人工林凋落物特性存在明显差异。凋落物各组分大量养分含量表现为N>Ca>Mg>K>P,微量养分表现为Fe>Mn>Zn>Cu,各组分养分浓度具有明显的月动态规律,以某一月份的养分浓度代表年平均浓度来计算凋落物年养分归还量会产生过高或过低的估计。以凋落物形式归还林地的养分量均表现为随发育阶段的增加而增加,大量养分归还量表现为过熟林(146.61kg·hm-2·a-1)>成龄林(131.46kg·hm-2·a-1)>幼龄林(96.63kg·hm-2·a-1),微量养分归还量表现为过熟林(12082.46g·hm-2·a-1)>成龄林(9087.19g·hm-2·a-1)>幼龄林(7796.22g·hm-2·a-1);大量元素养分年归还量总体表现为N>Ca>K或Mg>P,微量养分表现为Fe>Mn>Zn>Cu;各组养分归还量均表现为落叶或其它>落枝>落花果。杉木凋落物养分归还量具有明显的月动态特征,N归还量与凋落量月动态模式相似,幼龄林与成龄林表现为双峰型(5月与8月),过熟林表现为三峰型(1-2月、4-5月和8月)；P与K归还量的月动态表现三峰型,峰值出现的时间因发育阶段不同而不同；Ca与Mg归还量为单峰型(8月)；Fe养分归还量为多峰型,2、5、6、8均出现归还高峰；幼龄林与成龄林杉木Mn归还量月动态为单峰型(8月),过熟林为双峰型(5和8月)；三种发育阶段Cu, Zn归还量为双峰型(5、8月)。
相关分析表明,杉木林地内微量养分归还量与土壤养分及有效性关系更密切,微量养分元素可能与其它离子发生交互作用,促进土壤养分的转移及释放；杉木凋落物中N含量越高、C：N越低、纤维素和木质素含量越低,越有利于土壤有机质的矿化。
3、杉木人工林凋落物养分转移率较高,其中杉木枯枝和枯叶的N、P、K, Cu、Ca、Mg均存在不同程度的转移,而Fe、Mn、Zn均不发生养分转移。落叶的养分转移率表现为K或P>Cu>N>Ca>Mg,落枝的养分转移率表现为P>Cu>K>N或Ca> Mg,其中杉木落叶P转移率最高,不同发育阶段表现为成熟林(66.0%)>过熟林(65.4%)>幼龄林(60.9%),杉木落枝P转移率更高(71.0%-73.8%)；落枝叶N的转移率在30%-40%之间,落枝叶K的转移率在47%-70%之间；杉木落枝叶Ca, Mg转移率分别为4%-40%和4%-20%。养分转移率季节动态变化较大,表现为落枝叶N、P的转移率为双峰型(4月和10月),7月最低；落枝叶K的转移率为双峰型(4月和7月)；落枝叶Cu的转移率表同为4月最低(16.0%-24.7%),至7月开始升高,10月与1月仍保持较高的养分转移水平。影响杉木凋落物养分转移的各因素中,养分转移与新鲜和凋落组织的养分浓度关系比与受土壤影响更为密切,但养分转移并不仅仅是由高浓度向低浓度的简单迁移过程,而是与自身养分代谢有关的复杂过程。
杉木凋落物养分转移量(N、P、K、Ca、Mg、Cu6种元素)分别为25.51、39.03和47.25kg·hm-2·a-1,随发育阶段的增加而增加。其中落叶转移量为18.94-37.61kg·hm-2·a-1,落枝为5.57-10.09kg·hm-2·a-1。与归还量相比,养分转移量相当可观,不同发育阶段杉木人工林枝叶养分转移量分别占归还量的71.3%、57.9%和64.9%。各养分转移量排序为N>K>Ca>Mg>P>Cu。杉木人工林落枝叶养分转移量具有明显季节动态。三种杉木林分落叶N、P转移量高峰均出现在夏季(3.27-7.11kg·hm-2和0.21-0.59kg·hm-2);K转移高峰出现在春季和夏季；Cu转移高峰出现在冬季和夏季。
4、不同发育阶段杉木人工林细根生物量及时空分布差异明显。细根现存总量表现为成龄林(3.49t·hm-2)>过熟林(3.07t·hm-2)>幼龄林(2.80t·hm-2)；活细根现存量表现为成龄林(2.73t·hm-2)>幼龄林(2.49t·hm-2)>过熟林(2.13t·hm-2),表现为随发育阶段增加呈“∧”趋势。杉木细根垂直分布具有表聚性,细根现存总量(0-20cm)占40.4%-46.3%,活细根现存量(0-20cm)占47.4%-54.4%,但死细根垂直异质性不明显。杉木细根现存量的季节动态表明,细根现存总量表现为4月份出现第一次峰值,随后下降,7月份较低,10月份有所回升,出现次高峰；杉木活细根现存量1月份与4月份均较低,到7月份开始上升,10月份活细根现存量也较高；杉木死细根现存量月动态变化与活细根相反,表现为1、4月份较高,到7月份开始显著下降至最低,10月份略有回升。
不同发育阶段杉木人工林细根养分循环特征存在一定差异。幼龄林、成龄林和过熟林大量养分现存量分别为46.84、47.79和45.29kg·hm-2,微量养分现存量分别为1197.61、1720.49和1726.35g·hm-2；活细根养分现存量随发育阶段的增加而减小(大量养分：42.12、36.99和32.04kg·hm-2),死细根养分现存量与活细根相反,表现为随发育阶段的增加而增加(大量养分：4.74、10.81和13.24kg·hm-2(p<0.05)；微量养分：162.63、448.57和588.50g·hm-2(p<0.05))。大量养分现存量表现为N>K或Ca>Mg.P,微量养分现存量表现为Fe>Mn.Cu或Zn。细根养分的空间分布表明：杉木活细根N、P、K、Mg、Fe、Cu、Zn元素养分浓度随径级的增加而减小,Ca养分浓度随径级的增加而增加；死细根N、P、K养分浓度随径级的增加而减小,其它元素不同发育阶段表现规律不一；活细根N、P、K、Fe、Cu、Zn养分浓度随序级的增加而减小,死细根N、P、K养分浓度随序级的增加而减小,Ca和Mg随序级增加而增加。
不同发育阶段杉木人工林细根养分内循环研究表明：杉木细根K、Ca、Mg、Cu存在养分内循环,杉木细根N、P、Fe不发生养分内循环;各元素细根养分转移率总体表现为,各发育阶段细根养分转移率表现为幼龄林>过熟林>成龄林。杉木细根养分转移率与径级、序级相关性分析表明：K、Mg和Zn发生养分内循环的径级部位主要在0-0.5mm或0.5-1mm处,根序部位主要在1级或2级处。
Chinese fir (Cunninghamia lanceolata) is one of the most important native fast-growing evergreen coniferous species in southern China. Due to an increasing demand for timber, monoculture Chinese fir plantations were widely planted, however concerns have been expressed about the declining timber yield and soil fertility degradation on successive rotations of Chinese fir plantations. At present, the fundamental reasons contributing to the long-term problems are not clear. As a result, how to further reveal inter-mechanism of declining timber yield of Chinese fir plantations has become the major task in current forestry career.
The decline in Chinese fir productivity has been attributed to a combination of factors including site preparation methods, management intensity and silvicultural practices. However the role of the physiological characteristics of the species and nutrient cycling utilization in contributing to the decline in productivity is yet to be explored. Litterfall and fine root are important components of forest biogeochemical and nutrient cycling. Nutrient return and turnover through above-and belowground litterfall is an important pathway for self-fertilization in forests, which is a critical way to sustain the long-term productivity of Chinese-fir plantations. Although litterfall characteristics, fine root turnover and nutrient cycling utilization regarding to Chinese fir plantations has been studied, nutrient translocation and utilization efficiency was seldom studied and which has important theoretical and practical significance to understand the mechanism of productivity decline in Chinese fir plantations.
In view of this, this study focus on the problems in the nutrient cycling in Chinese fir plantations, from the aspect of the nutrient retranslocation and utilization, aboveground litterfall production, fine roots production, internal nutrient cycling in senescent leaf and branches and fine roots and nutrient distribution and fluxes were investigated in three monospecific Chinese-fir plantations at different developmental stages (10-,22-and34years) using the chronosequence method in Sanming city, Fujian province. The study will have important theoretical significance to understand the mechanism of productivity decline, also will have an important practical value to improve nutrient utilization and maximize forest productivity. The main results are as follows:
1. The biomass, productivity and nutrient utilization characteristic varied in different-stages Chinese fir plantations. Total biomass of the stands increased with the stage in the order of38.11,104.03and138.24t·hm-2respectively for the young, mature and over-mature stands. The total productivity also increased with stand stage, which were4.80,6.97and8.11t·hm-2·a-1for the10-,22, and34-year old stands respectively. Nutrient utilization efficiency in different developmental-staged Chinese fir plantation increased with the stage increasing. Producing dry matter per ton required macronutrient (N, P, K, Ca, Mg) of18.58,13.66and11.82kg respectively in young, mature and over-mature stage. Producing dry matter per ton required micronutrients (Fe, Mn, Cu, Zn) of0.608,0.490and0.490kg respectively in young, mature and over-mature stage. Nutrient utilization efficiency of macronutrient was in the order of P>Mg>Ca>K>N, and the micronutrient was in the order of Cu>Zn>Fe or Mn.
In conclusion, the young stage of Chinese-fir plantation was characterized by high nutrient uptake from the soil, low nutrient return, short turnover time and low nutrient utilization rate. In contrast, the mature stand circulation coefficient, turnover time, and nutrient utilization rate all increased, which indicates a higher nutrient uptake from soil as well as higher nutrient return and higher nutrient depletion. However, the turnover time in the over-mature stage stand was high and was characterized by lower uptake nutrient from the soil, higher nutrient return and utilization. This implies that harvesting Chinese-fir plantation at young stage would lead to high nutrient loss, which may not be sustainable in maintaining the long-term productivity of the species. As a result, prolonging the rotation of Chinese fir plantations properly will have great significance of soil fertility recovery.
2. For litterfall production, the height and layout mode of litter trap had no significant effect on litterfall collection with the results of50cm>25cm>100cm and "random" mold>"×"mold>"+" mold. The area of trap had significant effect on litterfall collection (p<0.05), with the order of0.5m2>2m2>1m2. The percentage of total trap area accounting for samples had significant effects on litterfall collection (p<0.05) in the order of1.5%>0.75%>0.5%. Annual litterfall production in different developmental-staged stands varied temporally. On average, annual litterfall production was in the order of over-mature stage (507.84g·m-2·a-1)> mature stage (458.79g·m-2·a-1)>young stage (348.52g·m-2·a-1). Reduced major axis analysis showed a significant positive correlation between litterfall fragments and total production. The allometric index between needles, branches, flowers and cones, other components and total litterfall production was1.36,1.41,1.52and1.31respectively. The allometric indexes between needles and branches, needles and flowers, branches and flowers, needles and others, branches and others, flowers and others were0.91,0.82,0.84,1.19,1.19and1.32respectively. With the exception of other components, needles accounted for the largest percentage of litterfall production (32.9%-44.4%), followed by branches (15.2%-17.2%), and flower and cones accounting for the smallest percentage (2.4%-10.8%). Monthly dynamics of litterfall production of Chinese fir plantations was obvious with three peaks of April-May, August and December. During2012-2013, the young and mature stage plantations had two peaks (April-May and August) in litterfall, however, monthly dynamics in the over-mature stage plantation was three peaks (April-May, August and January-February) in2012, and was one peak (April) in2013. Needles and branches had similar monthly dynamics as they dropped together. Seasonal dynamics of litterfall was in the order of spring> summer> autumn> winter. Leaves fell in spring, branches and flowers fell in summer.
Litterfall characteristics varied among different-staged Chinese fir plantations. Macronutrient concentration in litter of Chinese fir plantations was in the order of N> Ca> Mg> K> P, and micronutrient concentration was in the order of Mn> Fe> Zn> Cu.. Nutrient concentration in different litterfall components has obvious monthly dynamics, and it will be higher or lower evaluation of annual nutrient return by using nutrient concentration in one month. Macronutrient return (N, P, K, Ca and Mg) through litterfall increased with stand age, in the order of over-mature stage (146.61±25.19kg·hm-2·a-1)>mature stage (131.46±24.36kg·hm-2·a-1)>young stage (96.63±14.57kg·hm-2·a-1) respectively. Micronutrient return (Fe, Mn, Cu, Zn) also increased with stand age, following the order of over-mature stage (12082.46g-hm"2)> mature stage (9087.19g·hm-2)>young stage (7796.22g·hm-2) plantations respectively. The order of annual nutrient return by litterfall was N> Ca> K or Mg> P and Fe> Mn> Zn> Cu. Nutrient return in different components was in the order of leaves or other components>branches> flowers and cone. Monthly dynamics of N nutrient return was similar to litterfall production recording two peaks in the young and mature plantations (May and August), and three peaks in the over-mature stand (January-February, April-May and August). Monthly dynamics of P and K return were there peaks, and monthly dynamics of Ca and Mg had one peak in August. Monthly dynamics of Fe return had muti-peaks (Februay, May, June and August). Meanwhile the monthly dynamics of Mn return in the young and mature stands was one peak (August), and two peaks in the over-mature plantation (May and August). Monthly dynamics of Cu and Zn return were two peaks (May and August).
Pearson correlation showed significant relationship between C, K, Fe, Cu and Zn returned and litterfall production. The micronutrient returned through litterfall had a significant relationship with soil nutrient availability. N and C concentration was significantly correlated with soil organic matter. On the other hand P, C:N, cellulose content and lignin content had significantly negative correlation with soil organic matter. It can be concluded that the higher N content and the lower C:N and cellulose and lignin content is more advantageous to the mineralization of soil organic matter.
3. Independent-T test showed that N, P, K, Cu, Ca and Mg nutrients in senesced components retranslocated to the fresh ones, and Fe, Mn and Zn nutrients did not retranslocate when the leaves and branches senesced. Nutrients translocation rate of senesced needles was in the order of K> P> Cu> N> Ca> Mg, and the order for branches was P>Cu>K>N>Ca> Mg. The P nutrient retranslocation rate in leaves was higher following the order of over-mature stage (66.0%)>mature stage (65.4%)>young stage (60.9%), and the P nutrient retranslocation rate in branches was highest to71.0%-73.8%. The nutrient retranslocation rates of N, K, Ca, and Mg in senesced branches and leaves were30%-40%,47%-70%,4%-40%and4%-20%respectively. Monthly dynamics of N, P, K and Cu in senescenced leaves and branches was similar. N and P retranslocation rate peaked in April and October. Cu retranslocation rate was lowest in April (16.0%-24.7%), and higher in other months. Pearson correlation analysis showed that there was no significant correlation between nutrient retranslocation rate and soil nutrient pool. However, nutrient retranslocation rate was more related to nutrient content in fresh and dead tissues, but not a result of simple migration progress from high concentration to low concentration.
The nutrient retranslocation of N, P, K, Ca, Mg and Cu in leaves and branches increased with age in order of25.51,39.03, and47.25kg·hm-2·a-1for young, mature and over-mature stands respectively. The nutrient retranslocation amounts were18.94-37.61kg·hm-2·a-1in leaves and5.57-10.09kg-hm'^a"1in branches. Nutrient translocation amount through leaves and branches was considerable compared to nutrient return through litterfall, accounting for71.3%,57.9%and64.9%in different-staged Chinese fir plantations. Nutrient rethanslocation amount was in the order of N>K>Ca>Mg>P>Cu. Moreover seasonal dynamics of N, P, K and Cu retranslocation was apparent. The highest N and P retranslocation was recorded in summer (3.27-7.11kg·hm-2and0.21-0.59kg·hm-2) and the lowest was in autumn (1.33-4.59kg·hm-1); the highest K retranslocation amount was in spring in mature and over-mature stands and summer in young stand; the highest Cu retranslocation amount was in winter and summer in young and mature stands and winter in over-mature stand.
4. Fine root production and distribution varied in different-staged Chinese fir plantations. The total root biomass production was in the order of mature stage (3.49t-hm'2)>over-mature stage (3.07t·hm-2)> young stage (2.80t·hm-2) stands. The production of live fine roots was in the order of mature stage (2.73t·hm-2)> young stage (2.49t·hm-2)> over-mature stage (2.13t·hm-2) stands. The vertical distribution of fine roots of Chinese fir was accumulated in surface soil layer (0-20cm). The fine roots biomass production in0-20cm accounted for40.4%-46.3%of the total fine roots, and47.4%-54.4%of live fine roots. There was seasonal variation in fine root biomass production. Total production of fine root biomass showed two peaks with dominant peak in April and minor-peak in October. The production of live fine roots was high in January and April, while the pattern of dead fine roots showed opposite trend with high biomass in January and April and low biomass recorded in July and August.
Fine root nutrient characteristics varied in different-aged Chinese fir plantations. The macronutrient amount (N, P, K, Ca and Mg) of total fine roots for the young, mature and over-mature Chinese-fir stands were46.84,47.79, and45.29kg-hm'2respectively.The micronutrient concentration (Fe, Mn, Cu, Zn) of total fine roots were1197.61,1720.49and1726.35g·hm-2for the young, mature and over-mature stands respectively. The nutrient amount in live fine roots decreased with the stage increasing, while the nutrient amount in dead find roots increased with the stage increasing. The order of nutrient accumulation was in the order of N>K or Ca>Mg>P and Fe>Mn>Cu or Zn. Pearson correlation analysis showed that the concentration of N, P, K, Mg, Fe, Cu and Zn concentration in live fine roots decreased with root diameter increasing, while concentration of Ca in live fine roots showed increased trend with root diameter increasing. The concentration of N、P、K in dead fine roots increased with diameter increasing, while the other nutrients in dead fine roots showed different pattern. The concentration of N, P, K, Fe, Cu and Zn in live fine roots decreased with root order increasing. The concentration of N, P and K in dead fine roots decreased with root order increasing, while the concentration of Ca and Mg in dead fine roots increased with root order increasing.
Independent-T test showed that K, Ca, Mg, Cu nutrients retranslocated from dead roots to live roots, and N, P and Fe did not existed retranslocation when fine root seneced in Chinese fir plantations. The nutrient retranslocation rate was in the order of Cu>Ca>Mg>Mg and in the order of young stage> over-mature stage> mature stage. Pearson correlation analysis showed that K, Mg and Zn nutrient retranslocation exsted in the diameter of0.5-1mm and in the first and second order of fine roots.
有鉴于此,本研究针对杉木人工林养分循环研究中存在的问题,从养分内循环和周转利用效率入手,在福建农林大学莘口教学林场杉木人工林定位观测站的林分内,选择不同发育阶段的杉木幼龄林、成龄林和过熟林作为研究对象,研究不同发育阶段杉木人工林凋落物数量、养分归还、养分转移以及细根生长和衰老过程的养分转移,分析了不同发育阶段杉木林养分周转和利用效率,对提高杉木人工林生态系统的养分利用率和生产力水平具有重要现实意义,为深入揭示杉木人工林地力衰退的内在机制提供理论依据。主要研究结果如下：
1、不同发育阶段杉木人工林生物量、生产力水平及养分利用效率存在一定差异。生物量与生产力水平均表现为随发育阶段的增加而增加,幼龄林、成龄林和过熟林乔木层生物量分别为38.11、104.03和138.24t·hm-2,生产力分别为4.80、6.97和8.11t·hm-2·a-1。养分利用效率表现为随发育阶段的增加而增加,具体表现为林全树生产1吨干物质需要的大量养分量分别为18.58、13.66和12.74kg,微量养分量分别为0.608、0.490和0.490kg；大量养分利用效率表现为P>Mg>Ca>K>N,微量养分利用效率表现为Cu>Zn>Fe或Mn。通过比较各养分循环参数指标,表明幼龄林杉木具有高吸收、低归还、周转时间短、养分利用率低的特点,成熟林杉木养分循环系数提高,养分周转期延长,具有高吸收、高归还、养分消耗大特点,过熟林养分周转期延长,具有低吸收、高归还、养分利用效率高的特点。杉木生长后期更多关注自身生长,通过提高养分利用效率减少对地力的压力,若在杉木幼龄林时进行收获,势必带走大量养分,影响整个生态系统的平衡,因此,适当延长杉木轮伐期,对恢复地力意义重大,
2、通过对凋落物收集方式的筛选研究,结果表明：不同布点高度对收集量的影响表现为50cm>25cm>100cm;不同布点方式表现为“随机”>“×”>“+”；不同收集器面积表现为0.5m2>2m2>1m2(p<0.05)；收集器总面积占标准地的比例表现为1.5%>0.75%>0.5%。2012-2013年均凋落总量表现为随发育阶段的增加而增加,即过熟林(507.84g·m-2·a-1)>成龄林(458.79g·m-2·a-1)>幼龄林(348.52g·m-2·a-1),但具有年季变化特征。杉木凋落物各组分之间及其与总量之间存在异速比例关系,可以通过异速生长方程估测各组分的凋落量。叶、枝、花果、其它组分与总凋落量异速指数分别为1.36、1.41、1.52和1.31；叶与枝、叶与花果、枝与花果凋落量异速指数均小于1,分别为0.91、0.82和0.84,叶与其它、枝与其它和花与其它凋落量异速指均大于1,分别为1.19、1.19和1.32。凋落物各组分所占比例不同,落叶所占比例最大(32.9%-44.4%),枝次之(15.2%-17.2%),花果所占比例最小(2.4%-10.8%)。杉木人工林凋落物产量具有明显的季节节律,总体而言,凋落量出现在4-5月、8月和12月,针叶与枝凋落量月动态与总凋落量变化节律相似;季节变化表现为春季>夏季>秋季>冬季,叶主要集中在春季凋落,枝、花果凋落主要集中在夏季凋落。
不同发育阶段杉木人工林凋落物特性存在明显差异。凋落物各组分大量养分含量表现为N>Ca>Mg>K>P,微量养分表现为Fe>Mn>Zn>Cu,各组分养分浓度具有明显的月动态规律,以某一月份的养分浓度代表年平均浓度来计算凋落物年养分归还量会产生过高或过低的估计。以凋落物形式归还林地的养分量均表现为随发育阶段的增加而增加,大量养分归还量表现为过熟林(146.61kg·hm-2·a-1)>成龄林(131.46kg·hm-2·a-1)>幼龄林(96.63kg·hm-2·a-1),微量养分归还量表现为过熟林(12082.46g·hm-2·a-1)>成龄林(9087.19g·hm-2·a-1)>幼龄林(7796.22g·hm-2·a-1);大量元素养分年归还量总体表现为N>Ca>K或Mg>P,微量养分表现为Fe>Mn>Zn>Cu;各组养分归还量均表现为落叶或其它>落枝>落花果。杉木凋落物养分归还量具有明显的月动态特征,N归还量与凋落量月动态模式相似,幼龄林与成龄林表现为双峰型(5月与8月),过熟林表现为三峰型(1-2月、4-5月和8月)；P与K归还量的月动态表现三峰型,峰值出现的时间因发育阶段不同而不同；Ca与Mg归还量为单峰型(8月)；Fe养分归还量为多峰型,2、5、6、8均出现归还高峰；幼龄林与成龄林杉木Mn归还量月动态为单峰型(8月),过熟林为双峰型(5和8月)；三种发育阶段Cu, Zn归还量为双峰型(5、8月)。
相关分析表明,杉木林地内微量养分归还量与土壤养分及有效性关系更密切,微量养分元素可能与其它离子发生交互作用,促进土壤养分的转移及释放；杉木凋落物中N含量越高、C：N越低、纤维素和木质素含量越低,越有利于土壤有机质的矿化。
3、杉木人工林凋落物养分转移率较高,其中杉木枯枝和枯叶的N、P、K, Cu、Ca、Mg均存在不同程度的转移,而Fe、Mn、Zn均不发生养分转移。落叶的养分转移率表现为K或P>Cu>N>Ca>Mg,落枝的养分转移率表现为P>Cu>K>N或Ca> Mg,其中杉木落叶P转移率最高,不同发育阶段表现为成熟林(66.0%)>过熟林(65.4%)>幼龄林(60.9%),杉木落枝P转移率更高(71.0%-73.8%)；落枝叶N的转移率在30%-40%之间,落枝叶K的转移率在47%-70%之间；杉木落枝叶Ca, Mg转移率分别为4%-40%和4%-20%。养分转移率季节动态变化较大,表现为落枝叶N、P的转移率为双峰型(4月和10月),7月最低；落枝叶K的转移率为双峰型(4月和7月)；落枝叶Cu的转移率表同为4月最低(16.0%-24.7%),至7月开始升高,10月与1月仍保持较高的养分转移水平。影响杉木凋落物养分转移的各因素中,养分转移与新鲜和凋落组织的养分浓度关系比与受土壤影响更为密切,但养分转移并不仅仅是由高浓度向低浓度的简单迁移过程,而是与自身养分代谢有关的复杂过程。
杉木凋落物养分转移量(N、P、K、Ca、Mg、Cu6种元素)分别为25.51、39.03和47.25kg·hm-2·a-1,随发育阶段的增加而增加。其中落叶转移量为18.94-37.61kg·hm-2·a-1,落枝为5.57-10.09kg·hm-2·a-1。与归还量相比,养分转移量相当可观,不同发育阶段杉木人工林枝叶养分转移量分别占归还量的71.3%、57.9%和64.9%。各养分转移量排序为N>K>Ca>Mg>P>Cu。杉木人工林落枝叶养分转移量具有明显季节动态。三种杉木林分落叶N、P转移量高峰均出现在夏季(3.27-7.11kg·hm-2和0.21-0.59kg·hm-2);K转移高峰出现在春季和夏季；Cu转移高峰出现在冬季和夏季。
4、不同发育阶段杉木人工林细根生物量及时空分布差异明显。细根现存总量表现为成龄林(3.49t·hm-2)>过熟林(3.07t·hm-2)>幼龄林(2.80t·hm-2)；活细根现存量表现为成龄林(2.73t·hm-2)>幼龄林(2.49t·hm-2)>过熟林(2.13t·hm-2),表现为随发育阶段增加呈“∧”趋势。杉木细根垂直分布具有表聚性,细根现存总量(0-20cm)占40.4%-46.3%,活细根现存量(0-20cm)占47.4%-54.4%,但死细根垂直异质性不明显。杉木细根现存量的季节动态表明,细根现存总量表现为4月份出现第一次峰值,随后下降,7月份较低,10月份有所回升,出现次高峰；杉木活细根现存量1月份与4月份均较低,到7月份开始上升,10月份活细根现存量也较高；杉木死细根现存量月动态变化与活细根相反,表现为1、4月份较高,到7月份开始显著下降至最低,10月份略有回升。
不同发育阶段杉木人工林细根养分循环特征存在一定差异。幼龄林、成龄林和过熟林大量养分现存量分别为46.84、47.79和45.29kg·hm-2,微量养分现存量分别为1197.61、1720.49和1726.35g·hm-2；活细根养分现存量随发育阶段的增加而减小(大量养分：42.12、36.99和32.04kg·hm-2),死细根养分现存量与活细根相反,表现为随发育阶段的增加而增加(大量养分：4.74、10.81和13.24kg·hm-2(p<0.05)；微量养分：162.63、448.57和588.50g·hm-2(p<0.05))。大量养分现存量表现为N>K或Ca>Mg.P,微量养分现存量表现为Fe>Mn.Cu或Zn。细根养分的空间分布表明：杉木活细根N、P、K、Mg、Fe、Cu、Zn元素养分浓度随径级的增加而减小,Ca养分浓度随径级的增加而增加；死细根N、P、K养分浓度随径级的增加而减小,其它元素不同发育阶段表现规律不一；活细根N、P、K、Fe、Cu、Zn养分浓度随序级的增加而减小,死细根N、P、K养分浓度随序级的增加而减小,Ca和Mg随序级增加而增加。
不同发育阶段杉木人工林细根养分内循环研究表明：杉木细根K、Ca、Mg、Cu存在养分内循环,杉木细根N、P、Fe不发生养分内循环;各元素细根养分转移率总体表现为,各发育阶段细根养分转移率表现为幼龄林>过熟林>成龄林。杉木细根养分转移率与径级、序级相关性分析表明：K、Mg和Zn发生养分内循环的径级部位主要在0-0.5mm或0.5-1mm处,根序部位主要在1级或2级处。
Chinese fir (Cunninghamia lanceolata) is one of the most important native fast-growing evergreen coniferous species in southern China. Due to an increasing demand for timber, monoculture Chinese fir plantations were widely planted, however concerns have been expressed about the declining timber yield and soil fertility degradation on successive rotations of Chinese fir plantations. At present, the fundamental reasons contributing to the long-term problems are not clear. As a result, how to further reveal inter-mechanism of declining timber yield of Chinese fir plantations has become the major task in current forestry career.
The decline in Chinese fir productivity has been attributed to a combination of factors including site preparation methods, management intensity and silvicultural practices. However the role of the physiological characteristics of the species and nutrient cycling utilization in contributing to the decline in productivity is yet to be explored. Litterfall and fine root are important components of forest biogeochemical and nutrient cycling. Nutrient return and turnover through above-and belowground litterfall is an important pathway for self-fertilization in forests, which is a critical way to sustain the long-term productivity of Chinese-fir plantations. Although litterfall characteristics, fine root turnover and nutrient cycling utilization regarding to Chinese fir plantations has been studied, nutrient translocation and utilization efficiency was seldom studied and which has important theoretical and practical significance to understand the mechanism of productivity decline in Chinese fir plantations.
In view of this, this study focus on the problems in the nutrient cycling in Chinese fir plantations, from the aspect of the nutrient retranslocation and utilization, aboveground litterfall production, fine roots production, internal nutrient cycling in senescent leaf and branches and fine roots and nutrient distribution and fluxes were investigated in three monospecific Chinese-fir plantations at different developmental stages (10-,22-and34years) using the chronosequence method in Sanming city, Fujian province. The study will have important theoretical significance to understand the mechanism of productivity decline, also will have an important practical value to improve nutrient utilization and maximize forest productivity. The main results are as follows:
1. The biomass, productivity and nutrient utilization characteristic varied in different-stages Chinese fir plantations. Total biomass of the stands increased with the stage in the order of38.11,104.03and138.24t·hm-2respectively for the young, mature and over-mature stands. The total productivity also increased with stand stage, which were4.80,6.97and8.11t·hm-2·a-1for the10-,22, and34-year old stands respectively. Nutrient utilization efficiency in different developmental-staged Chinese fir plantation increased with the stage increasing. Producing dry matter per ton required macronutrient (N, P, K, Ca, Mg) of18.58,13.66and11.82kg respectively in young, mature and over-mature stage. Producing dry matter per ton required micronutrients (Fe, Mn, Cu, Zn) of0.608,0.490and0.490kg respectively in young, mature and over-mature stage. Nutrient utilization efficiency of macronutrient was in the order of P>Mg>Ca>K>N, and the micronutrient was in the order of Cu>Zn>Fe or Mn.
In conclusion, the young stage of Chinese-fir plantation was characterized by high nutrient uptake from the soil, low nutrient return, short turnover time and low nutrient utilization rate. In contrast, the mature stand circulation coefficient, turnover time, and nutrient utilization rate all increased, which indicates a higher nutrient uptake from soil as well as higher nutrient return and higher nutrient depletion. However, the turnover time in the over-mature stage stand was high and was characterized by lower uptake nutrient from the soil, higher nutrient return and utilization. This implies that harvesting Chinese-fir plantation at young stage would lead to high nutrient loss, which may not be sustainable in maintaining the long-term productivity of the species. As a result, prolonging the rotation of Chinese fir plantations properly will have great significance of soil fertility recovery.
2. For litterfall production, the height and layout mode of litter trap had no significant effect on litterfall collection with the results of50cm>25cm>100cm and "random" mold>"×"mold>"+" mold. The area of trap had significant effect on litterfall collection (p<0.05), with the order of0.5m2>2m2>1m2. The percentage of total trap area accounting for samples had significant effects on litterfall collection (p<0.05) in the order of1.5%>0.75%>0.5%. Annual litterfall production in different developmental-staged stands varied temporally. On average, annual litterfall production was in the order of over-mature stage (507.84g·m-2·a-1)> mature stage (458.79g·m-2·a-1)>young stage (348.52g·m-2·a-1). Reduced major axis analysis showed a significant positive correlation between litterfall fragments and total production. The allometric index between needles, branches, flowers and cones, other components and total litterfall production was1.36,1.41,1.52and1.31respectively. The allometric indexes between needles and branches, needles and flowers, branches and flowers, needles and others, branches and others, flowers and others were0.91,0.82,0.84,1.19,1.19and1.32respectively. With the exception of other components, needles accounted for the largest percentage of litterfall production (32.9%-44.4%), followed by branches (15.2%-17.2%), and flower and cones accounting for the smallest percentage (2.4%-10.8%). Monthly dynamics of litterfall production of Chinese fir plantations was obvious with three peaks of April-May, August and December. During2012-2013, the young and mature stage plantations had two peaks (April-May and August) in litterfall, however, monthly dynamics in the over-mature stage plantation was three peaks (April-May, August and January-February) in2012, and was one peak (April) in2013. Needles and branches had similar monthly dynamics as they dropped together. Seasonal dynamics of litterfall was in the order of spring> summer> autumn> winter. Leaves fell in spring, branches and flowers fell in summer.
Litterfall characteristics varied among different-staged Chinese fir plantations. Macronutrient concentration in litter of Chinese fir plantations was in the order of N> Ca> Mg> K> P, and micronutrient concentration was in the order of Mn> Fe> Zn> Cu.. Nutrient concentration in different litterfall components has obvious monthly dynamics, and it will be higher or lower evaluation of annual nutrient return by using nutrient concentration in one month. Macronutrient return (N, P, K, Ca and Mg) through litterfall increased with stand age, in the order of over-mature stage (146.61±25.19kg·hm-2·a-1)>mature stage (131.46±24.36kg·hm-2·a-1)>young stage (96.63±14.57kg·hm-2·a-1) respectively. Micronutrient return (Fe, Mn, Cu, Zn) also increased with stand age, following the order of over-mature stage (12082.46g-hm"2)> mature stage (9087.19g·hm-2)>young stage (7796.22g·hm-2) plantations respectively. The order of annual nutrient return by litterfall was N> Ca> K or Mg> P and Fe> Mn> Zn> Cu. Nutrient return in different components was in the order of leaves or other components>branches> flowers and cone. Monthly dynamics of N nutrient return was similar to litterfall production recording two peaks in the young and mature plantations (May and August), and three peaks in the over-mature stand (January-February, April-May and August). Monthly dynamics of P and K return were there peaks, and monthly dynamics of Ca and Mg had one peak in August. Monthly dynamics of Fe return had muti-peaks (Februay, May, June and August). Meanwhile the monthly dynamics of Mn return in the young and mature stands was one peak (August), and two peaks in the over-mature plantation (May and August). Monthly dynamics of Cu and Zn return were two peaks (May and August).
Pearson correlation showed significant relationship between C, K, Fe, Cu and Zn returned and litterfall production. The micronutrient returned through litterfall had a significant relationship with soil nutrient availability. N and C concentration was significantly correlated with soil organic matter. On the other hand P, C:N, cellulose content and lignin content had significantly negative correlation with soil organic matter. It can be concluded that the higher N content and the lower C:N and cellulose and lignin content is more advantageous to the mineralization of soil organic matter.
3. Independent-T test showed that N, P, K, Cu, Ca and Mg nutrients in senesced components retranslocated to the fresh ones, and Fe, Mn and Zn nutrients did not retranslocate when the leaves and branches senesced. Nutrients translocation rate of senesced needles was in the order of K> P> Cu> N> Ca> Mg, and the order for branches was P>Cu>K>N>Ca> Mg. The P nutrient retranslocation rate in leaves was higher following the order of over-mature stage (66.0%)>mature stage (65.4%)>young stage (60.9%), and the P nutrient retranslocation rate in branches was highest to71.0%-73.8%. The nutrient retranslocation rates of N, K, Ca, and Mg in senesced branches and leaves were30%-40%,47%-70%,4%-40%and4%-20%respectively. Monthly dynamics of N, P, K and Cu in senescenced leaves and branches was similar. N and P retranslocation rate peaked in April and October. Cu retranslocation rate was lowest in April (16.0%-24.7%), and higher in other months. Pearson correlation analysis showed that there was no significant correlation between nutrient retranslocation rate and soil nutrient pool. However, nutrient retranslocation rate was more related to nutrient content in fresh and dead tissues, but not a result of simple migration progress from high concentration to low concentration.
The nutrient retranslocation of N, P, K, Ca, Mg and Cu in leaves and branches increased with age in order of25.51,39.03, and47.25kg·hm-2·a-1for young, mature and over-mature stands respectively. The nutrient retranslocation amounts were18.94-37.61kg·hm-2·a-1in leaves and5.57-10.09kg-hm'^a"1in branches. Nutrient translocation amount through leaves and branches was considerable compared to nutrient return through litterfall, accounting for71.3%,57.9%and64.9%in different-staged Chinese fir plantations. Nutrient rethanslocation amount was in the order of N>K>Ca>Mg>P>Cu. Moreover seasonal dynamics of N, P, K and Cu retranslocation was apparent. The highest N and P retranslocation was recorded in summer (3.27-7.11kg·hm-2and0.21-0.59kg·hm-2) and the lowest was in autumn (1.33-4.59kg·hm-1); the highest K retranslocation amount was in spring in mature and over-mature stands and summer in young stand; the highest Cu retranslocation amount was in winter and summer in young and mature stands and winter in over-mature stand.
4. Fine root production and distribution varied in different-staged Chinese fir plantations. The total root biomass production was in the order of mature stage (3.49t-hm'2)>over-mature stage (3.07t·hm-2)> young stage (2.80t·hm-2) stands. The production of live fine roots was in the order of mature stage (2.73t·hm-2)> young stage (2.49t·hm-2)> over-mature stage (2.13t·hm-2) stands. The vertical distribution of fine roots of Chinese fir was accumulated in surface soil layer (0-20cm). The fine roots biomass production in0-20cm accounted for40.4%-46.3%of the total fine roots, and47.4%-54.4%of live fine roots. There was seasonal variation in fine root biomass production. Total production of fine root biomass showed two peaks with dominant peak in April and minor-peak in October. The production of live fine roots was high in January and April, while the pattern of dead fine roots showed opposite trend with high biomass in January and April and low biomass recorded in July and August.
Fine root nutrient characteristics varied in different-aged Chinese fir plantations. The macronutrient amount (N, P, K, Ca and Mg) of total fine roots for the young, mature and over-mature Chinese-fir stands were46.84,47.79, and45.29kg-hm'2respectively.The micronutrient concentration (Fe, Mn, Cu, Zn) of total fine roots were1197.61,1720.49and1726.35g·hm-2for the young, mature and over-mature stands respectively. The nutrient amount in live fine roots decreased with the stage increasing, while the nutrient amount in dead find roots increased with the stage increasing. The order of nutrient accumulation was in the order of N>K or Ca>Mg>P and Fe>Mn>Cu or Zn. Pearson correlation analysis showed that the concentration of N, P, K, Mg, Fe, Cu and Zn concentration in live fine roots decreased with root diameter increasing, while concentration of Ca in live fine roots showed increased trend with root diameter increasing. The concentration of N、P、K in dead fine roots increased with diameter increasing, while the other nutrients in dead fine roots showed different pattern. The concentration of N, P, K, Fe, Cu and Zn in live fine roots decreased with root order increasing. The concentration of N, P and K in dead fine roots decreased with root order increasing, while the concentration of Ca and Mg in dead fine roots increased with root order increasing.
Independent-T test showed that K, Ca, Mg, Cu nutrients retranslocated from dead roots to live roots, and N, P and Fe did not existed retranslocation when fine root seneced in Chinese fir plantations. The nutrient retranslocation rate was in the order of Cu>Ca>Mg>Mg and in the order of young stage> over-mature stage> mature stage. Pearson correlation analysis showed that K, Mg and Zn nutrient retranslocation exsted in the diameter of0.5-1mm and in the first and second order of fine roots.
引文
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