土壤有机碳和外源添加碳对水稻土土体呼吸的影响

土体呼吸输出碳来源于土壤固有有机碳和外源添加碳,而以往关于不同施肥措施对水稻土碳排放的研究少有区分碳的来源。本试验利用一个长达30年的水稻土定位试验,在保证原有定位试验继续正常开展的前提下变更部分施肥处理,得到继续施用高量有机肥(HOM)、施用常量有机肥30年后改施高量有机肥(N-H)、继续施用常量有机肥(NOM)、施用化肥30年后改施常量有机肥(C-N)、施用高量有机肥30年后改施化肥(H-C)、施用常量有机肥30年后改施化肥(N-C)、继续施用化肥(CF)等7种施肥处理。通过观测早稻生长期间原有施肥和改施肥处理土体CO2排放通量(FCO2),研究不同后续施肥对水稻土FCO2的影响,以期探讨土壤原始有机碳和外源添加碳对土壤FCO2的影响。结果表明:7种不同施肥处理土体CO2平均排放通量(F珔CO2)分别为85.34、69.10、51.27、49.15、14.89、12.92和11.59 mg C.m-2.h-1;对施用无机肥料和常量有机肥料的土体而言,土壤本身有机碳含量对F珔CO2无显著影响,但对施用高量有机肥的土体而言,土壤本身的高有机碳含量会增强F珔CO2;CO2排放通量(Y)与添加外源碳量(x)之间符合指数方程:Y=13.33e1.719 x(R2=0.967,n=21),施入的外源有机碳对土体FCO2产生极显著影响;当季外源添加碳以CO2-C矿化分解释放的碳占其总碳量的14%左右,且该分解率受土壤有机碳含量和有机物料添加量的影响较小。     

不同施肥措施下亚热带稻田土壤碳、氮演变特征及其耦合关系

在湖南省稻田生态系统长期定位监测点研究了不同施肥措施下稻田土壤碳、氮演变及其耦合特征.结果表明：1986—2003年,无肥处理（对照,CK）稻田土壤有机碳和全氮含量略呈下降趋势;化肥（NPK）处理有机碳和全氮含量基本保持稳定,而有机 无机肥配施处理有机碳和全氮含量均呈增加趋势.与对照相比,化肥处理的土壤有机碳和全氮含量分别提高13%和18%,低量有机肥（LOM）处理分别提高54%和45%,高量有机肥（HOM）处理分别提高89%和67%.统计分析表明,土壤有机碳和全氮含量呈显著正相关（P<0.01）.稻田土壤C/N为8.5～12.9,多数分布在10左右.研究表明,有机-无机肥配合施用能在一定程度上促进稻田土壤碳、氮的固定与积累;稻田土壤碳、氮具有较好的耦合关系.     

长期不同施肥下太湖地区黄泥土表土微生物碳氮量及基因多样性变化

农业管理措施影响下土壤微生物群落结构的变化是农业土壤质量研究的前沿问题。运用化学分析方法和 PCR- DGGE技术从土壤微生物碳氮量及基因多样性角度研究了长期不同施肥措施下太湖地区代表性水稻土 -黄泥土的表土微生物活性与分子多样性的变化。结果表明 ,施用化肥以及化肥和有机肥配施在提高土壤有机碳含量的同时 ,不仅提高了水稻土的微生物碳氮量 ,而且改变了微生物的群落结构 ;与长期单施化肥相比 ,长期化肥配施有机肥不仅显著提高了土壤微生物碳氮量 ,而且提高了土壤微生物的分子多样性 ;就土壤的微生物分子群落相似性来说 ,单施化肥下与未施肥下相近 ,而化肥配施秸秆下与化肥配施猪粪下接近 ,说明土壤的有机培肥对土壤微生物群落结构有重要影响。长期单施化肥下水稻产量的年际波动性显著大于化肥配施有机肥下 ,这进一步佐证了化肥配施有机肥显著促进了水稻土的生态系统初级生产力与较高的土壤生态系统稳定性。应用PCR- DGGE技术所揭示的微生物分子群落结构特点可以指示水稻土 10 a尺度的不同农业管理措施下的土壤质量变化     

不同地下水位和施肥管理对水稻土有机碳组分的影响

利用自1981年开始的红壤性水稻土长期定位试验,对耕层土壤不同有机碳组分进行分析,以探究不同地下水位(低水位80 cm、高水位20 cm)和施肥(高量有机肥、常量有机肥、化肥)管理对红壤性水稻土有机碳组分数量和分配的影响。结果表明:各处理土壤的颗粒有机碳(POC)含量为6.29~11.77 g·kg-1,高、低水位下,各施肥处理的POC含量均随着有机肥施用量的增加而增加,但高水位下POC增加的幅度更大;高、低水位下,有机肥处理的颗粒有机碳占总有机碳的比例(fPOC/TOC、oPOC/TOC)均显著高于化肥处理,且在低水位下这种差异尤为显著;土壤中各有机碳组分均与土壤总有机碳TOC和2 mm团聚体结合态有机碳含量存在极显著正相关;在高水位或低水位区,有机肥的施用均能大幅度提高土壤TOC和POC,且在高水位下有机碳的积累更为明显。     

不同培养温度下长期施肥水稻土的有机碳矿化特征

选择湖南省3个国家级稻田肥力变化长期定位监测点的土壤,分别在10、20和30℃进行室内恒温培养试验,研究温度和施肥对土壤有机碳矿化的影响及其与不同碳形态含量的关系.结果表明:在培养前期(0~13 d)土壤CO2产生速率较快,后期逐渐下降,其速率变化符合对数函数;升温促进了土壤有机碳矿化,各施肥处理土壤中,以秸秆还田和化肥配施有机肥处理土壤有机碳的累积矿化量较多;各施肥土壤的Q10值为1.01~1.53,与总有机碳、易氧化有机碳、胡敏酸碳和富里酸碳呈显著正相关;在10和20℃下CO2矿化量与微生物生物量呈显著的线性相关关系,而30℃下则无显著相关关系;CO2矿化量与不同碳形态含量和胡敏酸碳/富里酸碳呈显著的线性正相关;在10℃下矿化率与不同碳形态含量呈显著的线性负相关,在20和30℃下,矿化率与不同碳形态含量无明显相关关系.因此可以通过施用秸秆和有机肥增加对土壤碳的固定,减缓大气CO2浓度的升高,减少温室气体排放.     

稻田土壤氧化态有机碳组分变化及其与甲烷排放的关联性

稻田土壤有机碳是甲烷排放的关键底物之一,不同研究者由于采取的有机碳研究方法不同而得出稻田甲烷排放与土壤有机碳关系的结论不一.为明确影响稻田甲烷排放的土壤有机碳组分,设计了稻田施用不同外源有机碳(稻草还田、鸡粪和猪粪)的田间试验,对稻田甲烷排放和土壤有机碳组分的动态变化及其关联性进行监测和分析.结果表明,猪粪处理的甲烷排放与化肥处理无显著差异,而鸡粪和稻草2个处理的甲烷排放分别比化肥增加1.67倍(P＜0.05),2.69倍(P＜0.05);甲烷排放量与土壤易氧化有机碳含量显示相同顺序:稻草＞鸡粪＞猪粪＞化肥;通径分析表明,土壤易氧化有机碳组分1(被33 mmol/L KMnO4氧化的有机碳)与甲烷排放直接相关,其他有机碳组分仅通过组分1间接作用于水稻生育后期甲烷排放,且排放量较低.由此推断,易氧化有机碳组分1是甲烷排放的主要底物,通过有效措施降低肥源中易氧化态有机碳组分1是减排甲烷的关键技术之一.     

不同施肥对水稻土作物碳同化与土壤碳固定的影响——以太湖地区黄泥土肥料长期试验为例

肥料施用对农田土壤生产力及土壤碳循环的影响是农业与陆地生态系统碳循环及全球变化研究的重要科学问题。以太湖地区黄泥土的长期肥料定位试验为例,研究不同施肥处理对水稻土-作物系统作物碳同化及土壤碳固定的影响。所研究的肥料施用处理包括不施肥(NF)、单施化肥(CF)、化肥与秸秆配施(CFS)以及化肥与猪粪配施(CFM)4个处理,始于1987年,一直实行稻-油轮作,实行少耕。连续观测水稻和油菜的产量,并于2004年和2005年分别采集了土壤剖面样品和耕作层(0~5cm和5~15cm)土壤,测定土壤总有机碳含量。研究表明,不同施肥处理对水稻产量有显著影响,尤其以配施有机无机肥处理水稻产量显著最高且最为稳定,而对油菜产量的影响不明显。施肥显著提高了耕层土壤碳密度,而对全土碳密度没有显著影响。施肥处理的固碳速率介于0.1~0.4t/(hm2.a),配施有机肥处理显著高于单施化肥处理。相关分析表明,土壤固碳速率与作物根茬 有机肥源碳的总碳输入量呈显著的对数关系。这提示土壤有机碳积累主要与作物产量有关,而并非依变于有机肥源碳输入。因此,与作物产量直接关联的作物碳输入的增加是土壤中碳固定提高的重要途径。施入N素对水稻碳同化和土壤碳固定的效应均为化肥配施有机肥处理显著高于单施化肥处理,这揭示化肥配施有机肥是提高与稳定稻田生产力和促进土壤固碳和温室气体减排的双赢措施。当然,不同施肥下上述效应的差异可能与土壤-作物系统中碳分配和土壤生物碳利用的差异有关。     

长期施肥下红壤性水稻土有机碳储量变化特

研究了1982—2012年长期不同施肥下红壤性水稻土土壤有机碳含量变化、固碳趋势及外源碳输入对土壤固碳的贡献.结果表明： 施肥能提高土壤有机碳含量,连续30年不同施肥后,各施肥处理土壤有机碳含量趋于稳定,有机无机配施的土壤有机碳含量为21.02～21.24 g·kg^-1,增加速率为0.41～0.59 g·kg^-1·a^-1,单施化肥的土壤有机碳含量为15.48 g·kg^-1.各有机无机肥配施处理土壤的平均有机碳储量为43.61～48.43 t C·hm^-2,历年平均土壤有机碳储量显著大于单施化肥处理.土壤固碳速率与年均投入碳量呈显著指数正相关.本试验条件下,每年需要增加外源有机碳为0.12 t C·hm^-2才能维持土壤有机碳的平衡.     

不同施肥对水稻土作物碳同化与土壤碳固定的影响——以太湖地区黄泥土肥料长期试验为例

肥料施用对农田土壤生产力及土壤碳循环的影响是农业与陆地生态系统碳循环及全球变化研究的重要科学问题。以太湖地区黄泥土的长期肥料定位试验为例,研究不同施肥处理对水稻土-作物系统作物碳同化及土壤碳固定的影响。所研究的肥料施用处理包括不施肥(NF)、单施化肥(CF)、化肥与秸秆配施(CFS)以及化肥与猪粪配施(CFM) 4个处理,始于1987年,一直实行稻-油轮作,实行少耕。连续观测水稻和油菜的产量,并于2004年和2005年分别采集了土壤剖面样品和耕作层（0～5cm和5～15cm）土壤,测定土壤总有机碳含量。研究表明,不同施肥处理对水稻产量有显著影响,尤其以配施有机无机肥处理水稻产量显著最高且最为稳定,而对油菜产量的影响不明显。施肥显著提高了耕层土壤碳密度,而对全土碳密度没有显著影响。施肥处理的固碳速率介于0.1～0.4t/(hm^２•a), 配施有机肥处理显著高于单施化肥处理。相关分析表明,土壤固碳速率与作物根茬+有机肥源碳的总碳输入量呈显著的对数关系。这提示土壤有机碳积累主要与作物产量有关,而并非依变于有机肥源碳输入。因此,与作物产量直接关联的作物碳输入的增加是土壤中碳固定提高的重要途径。施入N素对水稻碳同化和土壤碳固定的效应均为化肥配施有机肥处理显著高于单施化肥处理,这揭示化肥配施有机肥是提高与稳定稻田生产力和促进土壤固碳和温室气体减排的双赢措施。当然,不同施肥下上述效应的差异可能与土壤-作物系统中碳分配和土壤生物碳利用的差异有关。     

长期施肥条件下黄土高原黑垆土作物产量与土壤碳氮的关系

通过设置在黄土高原黑垆土区的长期定位试验系统,研究了长期施肥条件下作物产量与土壤碳氮的互馈关系.试验设不施肥(CK)、单施氮肥(N)、氮磷配施(NP)、秸秆与氮磷配施(SNP)、施有机肥(M)和有机肥与氮磷配施(MNP)6个处理.结果表明: 与对照相比,长期平衡施用化肥、单施有机肥、化肥与有机肥配合施用和秸秆还田配施化肥显著增加了作物产量及其稳定性, NP、SNP、M、MNP处理玉米和小麦产量分别增加92%、97%、93%、141%和147%、164%、139%、214%.NP处理玉米和小麦年均产量与当地常规施肥作物产量相当且稳定,小麦-玉米轮作体系施肥量为N 90 kg·hm^-2、P₂O₅ 75 kg·hm^-2能够满足作物需要.秸秆还田与隔年施磷相配合的SNP处理与NP处理作物产量相似,且可减少磷肥施用量50%.平衡施用化肥、有机肥、化肥与有机肥配施和秸秆还田配施化肥均可显著增加土壤有机碳含量,而施用化肥对土壤全氮含量影响不明显,综合所有处理,土壤有机碳和全氮含量呈显著正相关.不同处理土壤有机碳固存率在15%～41%.SNP处理土壤有机碳累积投入量增加1 t·hm^-2,土壤有机碳含量增加0.06 g·kg^-1,而CK、N、NP、M和MNP处理的增幅在0.12～0.15 g·kg^-1.玉米和小麦产量都与土壤全氮含量呈显著正相关,玉米产量随土壤有机碳含量的增加而增加,但小麦产量随土壤有机碳含量的增加先快速增加后趋于平稳,拐点出现在6.8 g·kg^-1.长期平衡施用化肥、有机肥、有机肥与化肥配合施用及秸秆还田配施化肥可显著增加黄土高原黑垆土土壤有机碳和全氮含量、作物产量和根茬还田量,根茬还田量的增加又进一步增加了土壤有机碳和全氮含量,形成了相互促进的互馈关系.     

Carbon Sequestration in Bamboo Plantation Soil with Heavy Winter Organic Mulching Management

Carbon sequestration in soils is considered to be an important option for the mitigation of increasing atmospheric CO₂ concentrations as a result of climate change. High carbon accumulation was observed in Lei bamboo (Phyllostachys praecox) soils when using large amounts of organic material in a mulching technique. Soil samples were collected from Lei bamboo fields in a chronosequence. The composition and stability of soil organic carbon (SOC) in the bamboo soils was investigated by a combination of ¹³C CPMAS NMR analysis and with a decomposition incubation experiment in the laboratory. SOC content decreased in the first 5 years after planting of Lei bamboo from the original paddy soil and increased strongly subsequently. The stability of SOC after application of the winter mulch was higher as compared to the original paddy soil with no mulching, indicating that SOC can be stored effectively within Lei bamboo fields under intensive management.     

长期施肥对水稻土有机碳分布及化学结合形态的影响

采用湖南省4个23年连续施肥的稻田长期定位试验,研究了施肥对湖南省水稻土有机碳分布及化学键合形态的影响。试验设不施肥(CK)、化肥(NPK)、中量有机肥(MOM)和高量有机肥(HOM)4个处理。结果表明:在所有施肥处理中,水稳性团聚体均以0.25～1mm和2～5mm粒径含量最高,分别达全土总量的18%～43%和13%～48%。中、高量有机肥处理显著增加了>1mm大团聚体含量以及有机碳在大团聚体中的分配,其中0.25～1mm和1～2mm粒径团聚体中有机碳含量均略高于其余粒径组。与不施肥比较,钙结合态有机碳(Ca-SOC)占总有机碳的比例在2%～4%左右且随有机肥施用呈下降趋势,而铁铝结合态有机碳Fe(Al)-SOC占总有机碳的18%～33%呈上升趋势。有机肥施用条件下,有机碳在大团聚体中的分布的增加、Fe(Al)-SOC的提升以及Ca-SOC的降低意味着土壤有机碳物理和化学保护作用的增强,有利于稻田土壤有机碳的积累,是有机肥施用条件下保持稻田土壤较高固碳速率的重要原因。     

旱改水型农田整治对土壤碳排放的短期影响

灌溉农业可提升粮食生产潜力,已成为全球农业重要的发展方向,但此类土地利用转换势必影响旱作农田土壤的稳定性,尤其是碳循环。然而,旱改水整治过程中土壤碳通量变化及其与环境因子间的互馈机制尚不清楚。为此,采用大田模拟实验,连续7 d监测土壤碳通量变化,评估旱改水整治对土壤碳库组成及环境驱动的短期效应。结果表明：①旱地、水田的土壤碳通量和温度均呈昼高夜低的单峰型曲线,且碳通量与温度峰值出现于每日13：00前后,但水田土壤碳通量稍高于旱地。②旱改水后短期内土壤可溶性有机碳（DOC）、微生物量碳（MBC）、易氧化有机碳（EOC）、惰性有机碳（ROC）、总有机碳（TOC）和土壤碳库管理指数均呈减少趋势,其中土壤微生物量碳、易氧化有机碳降幅分别达28.55%、29.09%。③土壤含水量、微生物OTU数、碳库含量是影响碳通量速率变化的关键因子（P<0.05）,土壤温度、理化性状是制约土壤碳库的主控因子（P<0.05）。农业活动是重要的碳源之一,深入研究大范围旱改水诱发的碳排放问题可为低碳农业、气候减缓及其应对策略制定提供科学依据。     

Soil organic carbon budget and fertility variation of black soils in Northeast China

Black soils in Northeast China are characteristic of high soil organic carbon (SOC) density and were strongly influenced by human activities. Therefore, any change in SOC pool of these soils would not only impact the regional and global carbon cycle, but also affect the release and immobilization of nutrients. In this study, we reviewed the research progress on SOC storage, budget, variation, and fertility under different scenarios. The results showed that the organic carbon storage of black soils was 646.2 TgC and the most potential sequestration was 2887.8 g m⁻². According to the SOC budget, the net carbon emission of black soils was 1.3 TgC year⁻¹ under present soil management system. The simulation of CENTURY model showed that future climate change and elevated CO₂ concentration, especially the increase of precipitation, would increase SOC content. Furthermore, fertilization and cropping sequence obviously influenced SOC content, composition, and allocation among different soil particles. Long-term input of organic materials such as manure and straw renewed original SOC, improved soil structure and increased SOC accumulation. Besides, soil erosion preferred to transport soil particles with low density and fine size, decreased recalcitrant SOC fractions at erosion sites and increased activities of soil microorganism at deposition sites. After natural grasslands were converted into croplands, obvious variation of soil chemical nutrients, physical structure, and microbial activities had taken place in surface and subsurface soils, and represented a degrading trend to a certain degree. Our studies suggested that adopting optimal management such as conservation tillage in black soil region is an important approach to sequester atmospheric CO₂ and to slow greenhouse effects.     

不同施肥下中国旱地土壤有机碳变化特征——基于定位试验数据的Meta分析

搜集1994—2011年国内外有关中国旱地施肥处理的102个定位试验点的1146对田间试验数据,采用Meta-analysis方法定量分析了不同施肥条件下我国旱地耕层土壤有机碳(Soil Organic Carbon,SOC)的变化特征。结果表明,与对照(CK)相比,不同施肥措施均能显著提高耕层SOC含量,但不同施肥措施的效应不同。氮磷钾肥配施有机肥处理下SOC增速最大,为0.38 g kg-1a-1,单施磷肥处理增速最小,SOC增速仅为0.032 g kg-1a-1;添加有机肥的处理SOC增速远大于仅有无机化肥投入的施肥处理。不同施肥处理下SOC增速存在一定的空间分异特征且不同时期试验SOC相对变化速率也不相同,早期试验中SOC增速大于中、后期试验;不同种植制度对SOC变化速率的影响亦不同,有机肥的投入可以降低种植制度对SOC变化的影响。SOC积累与否及其幅度并不完全取决于初始SOC含量;随试验年限的增加,SOC增加速率呈降低趋势,仅采用短期试验(11 a)数据可能高估施肥措施下的固碳潜力。     

Response of soil carbon dioxide fluxes,soil organic carbon and microbial biomass carbon to biochar amendment: a meta‐analysis

Biochar as a carbon‐rich coproduct of pyrolyzing biomass, its amendment has been advocated as a potential strategy to soil carbon (C) sequestration. Updated data derived from 50 papers with 395 paired observations were reviewed using meta‐analysis procedures to examine responses of soil carbon dioxide (CO₂) fluxes, soil organic C (SOC), and soil microbial biomass C (MBC) contents to biochar amendment. When averaged across all studies, biochar amendment had no significant effect on soil CO₂ fluxes, but it significantly enhanced SOC content by 40% and MBC content by 18%. A positive response of soil CO₂ fluxes to biochar amendment was found in rice paddies, laboratory incubation studies, soils without vegetation, and unfertilized soils. Biochar amendment significantly increased soil MBC content in field studies, N‐fertilized soils, and soils with vegetation. Enhancement of SOC content following biochar amendment was the greatest in rice paddies among different land‐use types. Responses of soil CO₂ fluxes and MBC to biochar amendment varied with soil texture and pH. The use of biochar in combination with synthetic N fertilizer and waste compost fertilizer led to the greatest increases in soil CO₂ fluxes and MBC content, respectively. Both soil CO₂ fluxes and MBC responses to biochar amendment decreased with biochar application rate, pyrolysis temperature, or C/N ratio of biochar, while each increased SOC content enhancement. Among different biochar feedstock sources, positive responses of soil CO₂ fluxes and MBC were the highest for manure and crop residue feedstock sources, respectively. Soil CO₂ flux responses to biochar amendment decreased with pH of biochar, while biochars with pH of 8.1–9.0 had the greatest enhancement of SOC and MBC contents. Therefore, soil properties, land‐use type, agricultural practice, and biochar characteristics should be taken into account to assess the practical potential of biochar for mitigating climate change.     

Metagenomic and 14C tracing evidence for autotrophic microbial CO2 fixation in paddy soils

Autotrophic carbon dioxide (CO₂) fixation by microbes is ubiquitous in the environment and potentially contributes to the soil organic carbon (SOC) pool. However, the multiple autotrophic pathways of microbial carbon assimilation and fixation in paddy soils remain poorly characterized. In this study, we combine metagenomic analysis with ¹⁴C-labelling to investigate all known autotrophic pathways and CO₂ assimilation mechanisms in five typical paddy soils from southern China. Marker genes of six autotrophic pathways are detected in all soil samples, which are dominated by the cbbL genes (67%–82%) coding the ribulose-bisphosphate carboxylase large chain in the Calvin cycle. These marker genes are associated with a broad range of phototrophic and chemotrophic genera. Significant amounts of ¹⁴C-CO₂ are assimilated into SOC (74.3–175.8 mg ¹⁴C kg⁻¹) and microbial biomass (5.2–24.1 mg ¹⁴C kg⁻¹) after 45 days incubation, where more than 70% of ¹⁴C-SOC was concentrated in the relatively stable humin fractions. These results show that paddy soil microbes contain the genetic potential for autotrophic carbon fixation spreading over broad taxonomic ranges, and can incorporate atmospheric carbon into organic components, which ultimately contribute to the stable SOC pool.     

Influence of rhizodeposition under elevated CO2 on plant nutrition and soil organic matter

Atmospheric CO₂ concentrations can influence ecosystem carbon storage through net primary production (NPP), soil carbon storage, or both. In assessing the potential for carbon storage in terrestrial ecosystems under elevated CO₂, both NPP and processing of soil organic matter (SOM), as well as the multiple links between them, must be examined. Within this context, both the quantity and quality of carbon flux from roots to soil are important, since roots produce specialized compounds that enhance nutrient acquisition (affecting NPP), and since the flux of organic compounds from roots to soil fuels soil microbial activity (affecting processing of SOM).From the perspective of root physiology, a technique is described which uses genetically engineered bacteria to detect the distribution and amount of flux of particular compounds from single roots to non-sterile soils. Other experiments from several labs are noted which explore effects of elevated CO₂ on root acid phosphatase, phosphomonoesterase, and citrate production, all associated with phosphorus nutrition. From a soil perspective, effects of elevated CO₂ on the processing of SOM developed under a C4 grassland but planted with C3 California grassland species were examined under low (unamended) and high (amended with 20 g m^–2 NPK) nutrients; measurements of soil atmosphere 13C combined with soil respiration rates show that during vegetative growth in February, elevated CO₂ decreased respiration of carbon from C4 SOM in high nutrient soils but not in unamended soils.This emphasis on the impacts of carbon loss from roots on both NPP and SOM processing will be essential to understanding terrestrial ecosystem carbon storage under changing atmospheric CO₂ concentrations.Abbreviations SOM soil organic matter - NPP net primary productivity - NEP net ecosystem productivity - PNPP p-nitrophenyl phosphate