黔西北次生林优势树种叶片-凋落物-土壤连续体有机质碳稳定同位素特征

稳定性碳同位素自然丰度(δ13C)能够揭示生态系统长时间尺度的有机碳动态变化,阐明生态系统功能的变化特征。以黔西北次生林14个优势树种为研究对象,测定叶片、凋落物以及根区土壤有机碳含量和δ13C值,分析不同层次碳含量和δ13C丰度之间的相关性。结果表明:14个优势树种叶片碳含量为404.67—487.14 g/kg,总体为针叶树种较高、常绿灌木较低;δ13C值为-31.2‰—-27.1‰,随生活型的变化规律不明显。凋落物碳含量为414.62—561.31 g/kg,与叶片碳含量的变化规律较为一致;δ13C值为-31.5‰—-27.3‰,随树种生活型的变化特征也不明显。根区土壤碳含量为10.02—91.59 g/kg,δ13C值为-26.8‰—-22.5‰,碳含量以光皮桦、银白杨等落叶乔木较高。叶片、凋落物和根区土壤3个层次的碳含量与δ13C丰度之间均呈不显著相关,不同层次的δ13C丰度之间和碳含量之间均为正相关。研究结果有助于反映森林生态系统碳循环过程的关键信息,为森林植被恢复提供理论依据。     

长白山阔叶红松林演替序列植物-凋落物-土壤碳同位素特征

稳定碳同位素组成能精确指示生态系统碳循环过程,可以为深入研究森林演替进程对碳循环过程和固碳潜力的影响提供关键信息.利用稳定碳同位素技术对长白山阔叶红松林演替序列3种林分——中龄杨桦次生林、成熟杨桦次生林、阔叶红松林的叶片、树干、根系、凋落物和土壤δ¹³C值及碳、氮元素含量进行测定.结果表明: 各演替序列优势树种叶片δ¹³C从冠上到冠下均呈降低趋势;树干δ¹³C表现为树皮小于木质部;根系δ¹³C表现为细根小于粗根.阔叶红松林未分解凋落物δ¹³C小于半分解及全分解凋落物,次生林相反;土壤δ¹³C沿深度逐渐增加.总体上,δ¹³C值叶片<凋落物<根系<树干<土壤,说明植物各器官之间有明显的碳同位素分馏效应,且相同器官不同部位之间也存在差异;植物δ¹³C沿演替方向先减小后增加,土壤δ¹³C沿演替方向不断增加,且变化规律可以通过氮元素含量与碳同位素分馏效应的关系解释,说明长白山阔叶红松林演替过程优势树种和碳周转速率的变化影响了碳同位素分馏.     

半干旱沙丘与草甸典型植被类型区大气-叶片-凋落物-土壤连续体碳同位素特征

稳定碳同位素技术能够指示生态系统的物质循环与能量流动,根据生态系统碳转移动态,可以探明生态系统中碳循环过程和固碳能力。以科尔沁沙地半流动沙丘固沙植被差巴嘎蒿(Artemisia halodendron)、半流动半固定沙丘固沙植被小叶锦鸡儿(Caragana microphylla)和黄柳(Salix gordejevii Chang),以及在草甸地广泛分布的芦苇(Phragmites australis)与玉米(Zea mays Linn)5种典型植被为研究对象,分析了各植被群落冠层处大气、叶片、凋落物、土壤连续体的δ¹³C值和碳含量的分布特征及各组分间的关系。结果表明：沙丘植被冠层处大气CO₂浓度显著低于草甸植被,受控于土壤水分特征和植物生长特性。在逆境胁迫下,小叶锦鸡儿叶片水分利用效率最高,固碳耗水成本最低。叶片碳含量和δ¹³C值均受叶片生育期的影响,新叶片潜在碳蓄积能力更强,水分利用效率更高。叶片凋落物δ¹³C值在不同植被间存在显著差异,说明了植物功能性的驱动作用。随着土壤深度的增加,有机质分...     

基于稳定碳同位素分析宁夏典型林分土壤有机碳分布格局及影响因素

选择宁夏自北向南方向的贺兰山、罗山和六盘山(3座山的降水量呈现逐渐增加的趋势)共有的青海云杉林和油松林土壤作为研究对象,分析两种针叶林土壤有机碳特征[土壤有机碳(SOC)含量、稳定碳同位素组成(δ~(13)C)和有机碳分解速率(β值)]的变化情况,并采用冗余分析法分析土壤化学指标对土壤有机碳特征的影响作用,探究干旱半干旱区土壤稳定碳同位素特征,为揭示干旱区山地森林土壤碳动态变化规律提供依据。结果表明:(1)随样地降水量的增加,青海云杉和油松林地土壤SOC含量均逐渐增加,而土壤δ~(13)C和β值均逐渐减少,且在同等降水量条件下,青海云杉比油松更有利于土壤有机碳的累积。(2)青海云杉和油松林地土壤SOC含量均随土层深度的增加而下降,而其有机碳δ~(13)C均随土层深度的增加而增加。(3)宁夏贺兰山、罗山和六盘山青海云杉、油松森林土壤有机碳特征与其土壤化学指标间的冗余分析发现,碱解氮是影响土壤有机碳分布特征的主导因子。     

东北东部森林生态系统土壤碳贮量和碳通量

土壤碳是高纬度地区森林生态系统最大的碳库,是森林生态系统碳循环的极其重要组分。研究了东北东部典型的6种次生林生态系统(天然蒙古栎林、杨桦林、杂木林、硬阔叶林、红松人工林和落叶松人工林)的土壤碳动态,包括(1)量化土壤有机碳(SOC)含量、碳密度及周转时间,(2)比较不同森林生态系统的土壤表面CO2通量(RS)年通量差异,(3)建立RS年通量及其分量与SOC的量化关系。研究结果表明:阔叶天然次生林和针叶人工林的SOC含量变化范围分别为52.63~66.29 g.kg-1和42.15~49.15 g.kg-1;平均SOC密度分别为15.57和17.16 kg.m-2;平均SOC周转时间分别为32a和48a。各个生态系统的RS依次为杂木林951 gC.m-2.a-1、硬阔叶林892 gC.m-2.-a 1、杨桦林812 gC.m-2.-a 1、蒙古栎林678gC.m-2.-a 1、红松林596 gC.m-2.-a 1和落叶松林451 gC.m-2.a-1。RS年通量及其分量(土壤异养呼吸和自养呼吸)与SOC含量呈显著的正相关,但其相关程度因土层不同而异(R2=0.747~0.933)。同一生态系统中,SOC含量随土深增加而降低,而SOC密度和SOC周转时间随深度增加而增大。采用统一规范的研究方法,获取大量有代表性的森林生态系统土壤碳贮量和RS的实测数据,是减少区域尺度碳平衡研究中不确定性的不可缺少的研究内容。     

黄土高原植被演替对土壤碳库及δ~(13)C的影响

采集宁夏云雾山草原植被演替阶段的土壤和植物样品,测定土壤容重(BD)、土壤含水量(WC)、土壤有机碳(SOC)含量、以及土壤和植物优势种稳定碳同位素组成(δ13C),并比较了它们之间的差异,分析其反映的环境信息。结果表明:在表层0~10 cm深度,灌木地SOC含量为71.7 g·kg-1,显著高于草地的54.6 g·kg-1;灌木地0~20 cm深度SOC含量占1 m深SOC含量的58.1%,而草地在0~40 cm深度的SOC含量占1 m深的34.1%;短期植被演替(约10 a)对土壤剖面0~20 cm层的δ13C值有显著影响,其中对0~10 cm层δ13C影响最大;草地演替为灌木地后δ13C偏负1.4‰,30 cm以下土壤δ13C值对短期植被变化不敏感;通过δ13C在C3植被的短期演替过程中的响应关系发现,δ13C值作为土壤碳库更替和碳循环的研究工具有很好的辨识力;灌木的δ13C值从叶子到根系存在明显差异,逐渐偏正,变化为2.2‰,草地为1.9‰;通过对0~40 cm深度植物根系δ13C值的测定发现,灌木地由表层的-28.15‰变为-26.11‰,偏正了2.04‰,草地根系δ13C值从表层的-27.08‰变为-27.57‰,偏负了0.49‰。     

会仙喀斯特湿地三种典型植物叶片碳同位素(δ13C)特征及其指示意义

为探讨会仙喀斯特湿地不同生长环境下植物叶片碳(C)、氮(N)、磷(P)、稳定碳同位素(δ~(13)C)特征及其生态学指示意义,该文以挺水植物芦苇、浮水植物水葫芦和沉水植物金鱼藻为研究对象,分析了三种典型不同生活型植物叶片的δ~(13)C特征及种间和微生境的差异,并基于植物碳同位素与碳酸酐酶显著正相关的二端元模型,估算了植物利用光合作用固定的碳酸氢根离子(HCO3-)的碳量。结果表明:(1)三种植物叶片δ~(13)C的变化范围为-28.47‰～-21.69‰,平均值为-24.83‰,不同生活型植物间δ~(13)C存在差异,金鱼藻水葫芦芦苇。(2)植物δ~(13)C值与叶片C、N和P元素含量间呈显著正相关,与C/N、C/P和N/P呈显著负相关关系,与底泥的有机质、速效氮、总氮、速效磷和总磷含量呈显著正相关。(3)会仙喀斯特湿地三种不同生活型植物叶片N/P平均值为10.34,表现出植物受N、P共同影响的特征。(4)δ~(13)C的变化特征,揭示了三种水生植物可能通过增加磷利用效率来促进低水分利用率环境下的碳的合成,通过提高植物水分利用效率的策略来代偿较低的氮素利用效率。(5)芦苇光合作用固定HCO3-碳量为159.60 t·a~(-1)·km~(-2),水葫芦为10.80 t·a~(-1)·km~(-2),金鱼藻为9.24 t·a~(-1)·km~(-2),平均值为59.88 t·a~(-1)·km~(-2)。会仙喀斯特湿地植物的不同生活型、光合作用途径和生长微环境,是影响叶片δ~(13)C变化的主要因素。     

模拟增温对高寒草甸植物叶片碳氮及其同位素δ~(13)C和δ~(15)N含量的影响

基于国际冻原计划(ITEX)模拟增温效应对植物影响的研究方法,以高寒矮嵩草草甸4种植物(矮嵩草、垂穗披碱草、棘豆、麻花艽)为实验材料,设置大(OTC1)和小(OTC2)两类增温小室,测定了其叶片碳氮及其稳定性碳同位素(δ~(13)C、δ~(15)N)等指标在增温3年后的变化。结果表明:除矮嵩草在较小增温小室(OTC2)中C/N比值比对照降低了14.1%,其它物种C/N在两个增温处理下都有所增加,但差异均不显著(P0.05)。4种植物叶片δ~(13)C值在-24.12‰~-28.34‰之间,矮嵩草叶片δ~(13)C值随增温而升高,棘豆、麻花艽随增温而降低,且矮嵩草在OTC2的δ~(13)C值变化达到显著水平(P0.05)。矮嵩草和麻花艽的叶片δ~(15)N值在OTC1和OTC2中均比对照增加,且麻花艽增加较显著(P0.05)。垂穗披碱草在OTC1和OTC2的叶片δ~(15)N值比对照分别减少18.7%和26.9%,差异都不显著(P0.05);棘豆叶片δ~(15)N值在OTC2内比对照低11.0%(P0.05),在OTC1的内比对照高2.8%(P0.05)。可见,高寒矮嵩草草甸不同功能群植物物种碳氮含量及稳定性碳氮同位素含量对短期增温有不同的响应模式和规律。     

湖泊水生植物稳定碳同位素分馏机制与应用研究进展

水生植物是维持湖泊生态系统健康发展的重要基础与支撑。与陆地植物相比,水生植物稳定碳同位素(δ~(13)C)的分馏过程较复杂,为系统的了解水生植物δ~(13)C的分馏过程及其在湖泊现代生态系统和古环境研究中的应用,调研了国内外相关研究进展。总结已有研究表明,水生植物的δ~(13)C主要与其吸收利用的碳源和水体环境要素等密切相关,影响因素较复杂,而且目前在利用水生植物的δ~(13)C研究不同时间尺度上湖泊环境变化时所选用的载体存在差异。植物不同组分样品的选择对解释水生植物δ~(13)C与环境之间的关系至关重要。因此,以太湖典型水生植物为研究实例,探讨了水生植物不同组分稳定碳同位素的分布特征及其对环境因子的响应差异,结果显示水生植物不同组分的δ~(13)C存在一定的差异,而且对环境的响应也不同,未来在利用水生植物δ~(13)C研究湖泊环境变化时对于水生植物样品组分的选择应谨慎考虑。     

13C脉冲标记法：不同生育期水稻光合碳在植物-土壤系统中的分配

定量生育期内植物光合碳在植物组织-土壤的分配规律,对于理解全球碳循环有着重要意义。采用~(13)C-CO_2脉冲标记结合室内培养,通过元素分析仪-稳定同位素联用(Flash HT-IRMS)分析植物各部分及土壤δ~(13)C值,比较了不同生育期下水稻光合碳在不同组织间的分配规律,并量化了水稻光合碳向土壤碳库的转移。结果表明:(1)水稻地上部和根系干物质量随水稻生育期的增加而呈现递增趋势,不同的生育期表现为:分蘖期拔节期抽穗期扬花期成熟期。而整个生育期的根冠比为0.2—0.4,分蘖期的根冠比最高,随着水稻生育期的增加而递减,到抽穗期以后根冠比稳定在0.2左右。(2)脉冲标记6h后,水稻地上部和地下部(根系)的δ~(13)C值在-25.52‰—-28.33‰,不同器官的δ~(13)C值存在明显分馏效应,且趋势基本一致,即茎杆(籽粒)叶片(根系);这种由于水稻生育期特性导致的各器官碳同位素分馏的现象,可用于指示不同生育期下水稻光合碳的分配和去向。(3)不同生育期~(13)C-光合碳在植物-土壤系统的分配规律不同,生长前期光合碳向根系及土壤中分配的比例高,具有较强的碳汇能力,而随生育期光合碳在根系及土壤中的分配比例呈下降趋势,但积累量不断增加。(4)不同生育期~(13)C光合碳在水稻-土壤系统中的分配比例差异明显。水稻分蘖期有近30%光合碳用于根系建成并部分通过根系分泌物进入土壤有机碳库(10%),而到成熟期则向籽粒中分配较多,而且光合碳在土壤中的分配比例也随生育期呈下降趋势。研究结果对稻田土壤有机碳循环过程和调控机制的揭示具有重要的理论意义。     

Scope for use of stable carbon isotopes in discerning the incorporation of forest detritus into aquatic foodwebs

Stable isotope analysis of carbon has been proposed as a means for discerning the incorporation of terrestrial forest detritus into aquatic foodwebs, and as such, has the potential to be used as a biomonitor of the aquatic effects of riparian deforestation. A synthesis of ¹³C/¹²C data from the literature indicates, however, that the scope for successful use of carbon isotope analysis in separating allochthonous and autochthonous food provenance is much more limited than was once thought. This occurs due the overlap in carbon isotope ratios between terrestrial forest detritus and those of both lotic attached algae and lentic filamentous attached algae. Only within rockyshored, oligotrophic lakes without macrophytes, and forest-fringed estuaries and lagoons, where the carbon isotope ratios for attached algae and forest detritus are significantly different, is there any likelihood of discerning the incorporation of allochthonous carbon into aquatic foodwebs using ¹³C/¹²C values alone.     

Assimilate partitioning affects 13C fractionation of recently assimilated carbon in maize

Coupling ¹³C natural abundance and ¹⁴C pulse labelling enabled us to investigate the dependence of ¹³C fractionation on assimilate partitioning between shoots, roots, exudates, and CO₂ respired by maize roots. The amount of recently assimilated C in these four pools was controlled by three levels of nutrient supply: full nutrient supply (NS), 10 times diluted nutrient supply (DNS), and deionised water (DW). After pulse labelling of maize shoots in a ¹⁴CO₂ atmosphere, ¹⁴C was traced to determine the amounts of recently assimilated C in the four pools and the δ¹³C values of the four pools were measured. Increasing amounts of recently assimilated C in the roots (from 8% to 10% of recovered ¹⁴C in NS and DNS treatments) led to a 0.3‰ ¹³C enrichment from NS to DNS treatments. A further increase of C allocation in the roots (from 10% to 13% of recovered ¹⁴C in DNS and DW treatments) resulted in an additional enrichment of the roots from DNS to DW treatments by 0.3‰. These findings support the hypothesis that ¹³C enrichment in a pool increases with an increasing amount of C transferred into that pool. δ¹³C of CO₂ evolved by root respiration was similar to that of the roots in DNS and DW treatments. However, if the amount of recently assimilated C in root respiration was reduced (NS treatment), the respired CO₂ became 0.7‰ ¹³C depleted compared to roots. Increasing amounts of recently assimilated C in the CO₂ from NS via DNS to DW treatments resulted in a 1.6‰ δ¹³C increase of root respired CO₂ from NS to DW treatments. Thus, for both pools, i.e. roots and root respiration, increasing amounts of recently assimilated C in the pool led to a δ¹³C increase. In DW and DNS plants there was no ¹³C fractionation between roots and exudates. However, high nutrient supply decreased the amount of recently assimilated C in exudates compared to the other two treatments and led to a 5.3‰ ¹³C enrichment in exudates compared to roots. We conclude that ¹³C discrimination between plant pools and within processes such as exudation and root respiration is not constant but strongly depends on the amount of C in the respective pool and on partitioning of recently assimilated C between plant pools. Section Editor: H. Lambers     

The role of the external mycelial network of vesicular-arbuscular mycorrhizal fungi: a study of carbon transfer between plants interconnected by a common mycelium

The transfer of ¹⁴C from Lolium perenne (the donor) to Plantago lanceolata (the receiver), mediated by vesicular-arbuscular (VA) mycorrhizal fungi, was examined when the two species were grown together or separately. The VA mycorrhizal infection led to a significant increase, relative to that in uninfected plants, in the ¹⁴C transferred from donor to receiver plants, not only when the roots of the two plants were growing in intimate mixture, but also when they were separated by a root-free zone of 2.33 cm. The majority of isotope transfer between the two plant species was along the direct pathway via VA mycelium.     

Comparisons of carbon isotope discrimination in populations of aridland plant species differing in lifespan

Summary Carbon isotope discrimination () was compared between populations of dominant perennial plant species, differing in life expectancy, in two deserts with contrasting vegetation types. In both deserts, plants of the shorter-lived species showed significantly higher and greater intrapopulation variance in this character compared to the long-lived species. These results indicate underlying differences in gas-exchange physiology, and suggest a positive correlation between water-use efficiency and lifespan in desert plants. Differences in variance for this character may reflect greater microenvironmental variation experienced by shorter-lived plants and/or different forms of selection acting on water-use traits. Spatial distributions were significantly clustered for the shorter-lived species and significantly uniform for the long-lived species, indicating that competition has been important in the development of the long-lived populations. The long-lived Larrea tridentata showed a significant, negative correlation between and Thiessen polygon area, suggesting a positive relationship between water-use efficiency and longevity within this species. This relationship was weakly supported in the other warm desert species, Encelia farinosa, but was not observed within populations of the cold desert species, Gutierrezia microcephala and Coleogyne ramosissima. These results suggest that reflects key aspects of plant metabolism related to lifespan; these differences may ultimately influence interactions among desert plants and the structure of desert plant communities.     

Leaf carbon isotope and mineral composition in subtropical plants along an irradiance cline

Summary Leaf carbon isotope ratios and leaf mineral composition (Ca, K, Mg, Mn, N, and P) were measured on the dominant species along an irradiance cline in a subtropical monsoon forest of southern China. This irradiance cline resulted from disturbance caused by fuel-harvesting. Leaf carbon isotope ratios increased from undisturbed to disturbed sites for all species, indicating that leaf intercellular CO₂ concentrations decreased and leaf water use efficiencies increased along this cline. Nitrogen and magnesium levels were lower in leaves of species on the disturbed sites, but there were no clear patterns for calcium, potassium, phosphorus or manganese.C.I.W.D.P.B. Publication no 932     

Inhibition of trehalose breakdown increases new carbon partitioning into cellulosic biomass in Nicotiana tabacum

Validamycin A was used to inhibit in vivo trehalase activity in tobacco enabling the study of subsequent changes in new C partitioning into cellulosic biomass and lignin precursors. After 12-h exposure to treatment, plants were pulse labeled using radioactive ¹¹CO₂, and the partitioning of isotope was traced into [¹¹C]cellulose and [¹¹C]hemicellulose, as well as into [¹¹C]phenylalanine, the precursor for lignin. Over this time course of treatment, new carbon partitioning into hemicellulose and cellulose was increased, while new carbon partitioning into phenylalanine was decreased. This trend was accompanied by a decrease in phenylalanine ammonia-lyase activity. After 4 d of exposure to validamycin A, we also measured leaf protein content and key C and N metabolite pools. Extended treatment increased foliar cellulose and starch content, decreased sucrose, and total amino acid and nitrate content, and had no effect on total protein.     

Application of <Superscript>13</Superscript>C mineral carbon for assessment of the primary production of organic matter in aquatic environments

The possibility of measuring the rates of light and dark CO₂ assimilation using ¹³C carbonate was demonstrated on Lake Kichier (Marii El). The application of methods utilizing the stable ¹³C and the radioactive ¹⁴C isotopes resulted in comparable values of the rates of light and dark CO₂ fixation. Due to its absolute environmental safety, the method with ¹³C mineral carbon can be recommended as an alternative to radioisotope methods for qualitative measurements of CO₂ fixation rates in aquatic ecosystems.     

Allocation of carbon to shoots,roots, soil and rhizosphere respiration by barrel medic (Medicago truncatula) before and after defoliation

The allocation of carbon to shoots, roots, soil and rhizosphere respiration in barrel medic (Medicago truncatulaGaertn.) before and after defoliation was determined by growing plants in pots in a labelled atmosphere in a growth cabinet. Plants were grown in a ¹⁴CO₂-labelled atmosphere for 30 days, defoliated and then grown in a ¹³CO₂-labelled atmosphere for 19 days. Allocation of ¹⁴C-labelled C to shoots, roots, soil and rhizosphere respiration was determined before defoliation and the allocation of ¹⁴C and ¹³C was determined for the period after defoliation. Before defoliation, 38.4% of assimilated C was allocated below ground, whereas after defoliation it was 19.9%. Over the entire length of the experiment, the proportion of net assimilated carbon allocated below ground was 30.3%. Of this, 46% was found in the roots, 22% in the soil and 32% was recovered as rhizosphere respiration. There was no net translocation of assimilate from roots to new shoot tissue after defoliation, indicating that all new shoot growth arose from above-ground stores and newly assimilated carbon. The rate of rhizosphere respiration decreased immediately after defoliation, but after 8 days, was at comparable levels to those before defoliation. It was not until 14 days after defoliation that the amount of respiration from newly assimilated C (¹³C) exceeded that of C assimilated before defoliation (¹⁴C). This revised version was published online in June 2006 with corrections to the Cover Date.     

Stable carbon and nitrogen isotopic discrimination in whole blood of red-throated ant tanagers Habia fuscicauda

Analysis of stable isotope ratios is increasingly used to reconstruct diets in passerine birds, but studies of diet–tissue isotopic discrimination for this avian group are scarce. We determined ¹⁵N and ¹³C diet–tissue discrimination factors on whole blood in the red-throated ant tanager (Habia fuscicauda), an insectivorous–frugivorous passerine. Birds were fed an isotopically uniform, semi-synthetic diet of dog puppy dry food, soy protein isolate, wheat germ, and other ingredients, during 92 days. Average (± SD) diet–tissue discrimination was 2.6 ± 0.2‰ for N and 2.2 ± 0.1‰ for C. Nitrogen diet-tissue discrimination was similar to the values found previously in other passerines fed animal protein and it can probably be used to accurately reconstruct protein dietary origin in passerines feeding on animal protein (e.g., insects). In the case of C, diet reconstruction might be affected by metabolic routing of dietary nutrients.     

Below-ground carbon distribution in barley (Hordeum vulgare L.) with and without nitrogen fertilization

The distribution of net assimilated C in barley (Hordeum vulgare L.) grown at two N-levels was determined in a growth chamber. The N-fertilization involved 0 and 3.61 mol N g^-1 dry soil. After growth for seven weeks in an atmosphere with continuously ¹⁴C-labelled CO₂, ¹⁴C was determined in shoots, roots, rhizosphere respiration and soil. At the low N-level, 32% of the net assimilated ¹⁴C was translocated below ground, whereas at the high N-level 27% was translocated below ground. The release of C from roots (root respiration, microbial respiration originating from decomposition of ¹⁴C-labelled root material and ¹⁴C remaining in soil) was greater with no N-supply (19% of net assimilated ¹⁴C) than in the treatment with N-supply (15%). Thus, the effect of N-supply on both translocation of assimilated ¹⁴C below ground and the release of ¹⁴C from growing roots was relatively small.