Gas exchanges and chlorophyll fluorescence of young coffee plants submitted to water and nitrogen stresses

Abstract

This research aims to evaluate the impact of nitrogen deprivation and water stress on gas exchange and chlorophyll fluorescence in young plants of five cultivars of Arabic coffee. A factorial experiment 5 (cultivars) × 3 (treatments: control without stress, water stress of ?1.5?MPa and stress of N – 0.0?mmol L^?1 N) was carried out in a complete randomized block design with three replicates. Before being submitted to the treatments, the plants were grown in a greenhouse for 240?days, and then transferred to a growth chamber under controlled conditions. Subsequently, after the experimental period of 96?h we measured photosynthetic rate (A), stomatal conductance to water vapor (g_s), transpiratory rate (E), internal and external carbon ratio (Ci/Ca), water use efficiency (A/E), electron transport rate (ETR), actual quantum yield of PS II electron transport (φFSII), and maximum photochemical efficiency of PS II (Fv/Fm). Water stress reduced A, g_s, E, A/E, ETR, φFSII, and Fv/Fm. The nitrogen deficiency reduced ETR, φFSII, and Fv/Fm. Under short-term water stress Catuaí Vermelho maintain the A values due to better stomatal control, reduced water lost by transpiration (E) and better water use efficiency A/E, while Mundo Novo and Acauã show lower damage to Fv/Fm. Short-term nitrogen stress has low impact on A of young plants of Coffea arabica cultivars with adequate N-nutrition.     

Effect of copper sprays oh the mineral nutrient content and growth of arabica coffee seedlings in Kenya

Abstract

Coffea arabica L. seedlings (cv. S.L.34) were sprayed with varying concentrations of cuprous oxide at 0.0, 0.25, 0.50 and 0.75% for 30 days. Significant increases in the total mean fresh and dry weight; rate of transpiration; stomatal apertures; the total leaf area and plant vigor were observed on the seedlings with increasing concentrations of Cu treatments. The content of N, K, Ca and Cu were significantly increased in the seedlings at Cu spray treatments of 0.50 and 0.75% It is possible that the growth promoting effects produced in coffee by spraying high concentrations of copper is partly nutritional. This is brought about by the catalysing effects of the absorbed Cu ions, by accelerating enzymatic activities within the plant and hence increasing nutrient uptake.     

Effect of soil drying on mucilage exudation and its water repellency: a new method to collect mucilage

Despite the importance of mucilage for soil–plant relations, little is known about the effect of soil drying on mucilage exudation. We introduce a method to collect mucilage from maize growing in wet and dry soils. Mucilage was collected from brace roots. The amount of mucilage exuded did not change with soil water content and transpiration rate. Mucilage exuded in dry soils had a higher degree of hydrophobicity, suggesting that the wetting properties of mucilage change in response to soil drying.     

分根区交替灌溉对玉米水分利用和土壤微生物量碳的影响

分根区交替灌溉由于创造了一个土壤水分分布不均匀的环境,从而影响土壤中微生物活性,作物水分和养分利用。为探明这种影响,该文通过盆栽试验,研究了在2种灌水水平(正常灌水W1,70%～80%田间持水率;轻度缺水W2,60%～70%田间持水率)和2种有机无机氮比例(100%无机氮,70%无机氮+30%有机氮)条件下,常规灌溉和不同生育期分根区交替灌溉(分别在苗期～灌浆初期、苗期～拔节期以及拔节期～抽雄期进行分根区交替灌溉(AI),即AI1、AI2和AI3)对玉米干物质积累、水分利用以及拔节期、抽雄期和灌浆初期土壤微生物量碳(MBC),可溶性碳(DOC)含量以及基础呼吸和诱导呼吸CO2释放量等的影响。结果表明,与常规灌溉相比,轻度缺水时,拔节期～抽雄期分根区交替灌溉总干物质质量增加23.2%～27.4%,水分利用效率提高23.3%～26.7%;相同施肥和灌水水平条件下,抽雄期时拔节期～抽雄期分根区交替灌溉土壤MBC增加,但是土壤诱导呼吸CO2释放量降低。与单施无机氮相比,有机、无机氮配施增加玉米干物质质量,在某些水分条件下(W1CI、W1AI1和W1AI2)还提高灌浆初期基础呼吸和诱导呼吸CO2释放量。因此,轻度缺水时拔节期～抽雄期进行分根区交替灌溉可以提高玉米总干物质质量、水分利用效率和微生物量碳。     

Perspectives from the Fritz‐Scheffer Awardee 2017. How mucilage affects soil hydraulic dynamics#

Mucilage is a hydrogel exuded at root tips, which can hold large amounts of water but turns hydrophobic once dried. It is very challenging to understand the interplay of these opposite mechanisms and to incorporated them into hydraulic soil models. My summary of experimental and modelling approaches and observations at various scales is meant to help improving soil–plant water dynamic models and may also be helpful when water dynamics need to be considered in biogeochemical rhizosphere models.     

Rate of soil‐aggregate formation under different organic matter amendments—a short‐term incubation experiment

To improve soil structure and take advantage of several accompanying ecological benefits, it is necessary to understand the underlying processes of aggregate dynamics in soils. Our objective was to quantify macroaggregate (> 250 μm) rebuilding in soils from loess (Haplic Luvisol) with different initial soil organic C (SOC) contents and different amendments of organic matter (OM) in a short term incubation experiment. Two soils differing in C content and sampled at 0–5 and 5–25 cm soil depths were incubated after macroaggregate destruction. The following treatments were applied: (1) control (without any addition), (2) OM₁ (addition of OM: preincubated wheat straw [< 10 mm, C : N 40.6] at a rate of 4.1 g C [kg soil]^–1), and (3) OM₂ (same as (2) at a rate of 8.2 g C [kg soil]^–1). Evolution of CO₂ released from the treatments was measured continuously, and contents of different water‐stable aggregate‐size classes (> 250 μm, 250–53 μm, < 53 μm), microbial biomass, and ergosterol were determined after 7 and 28 d of incubation. Highest microbial activity was observed in the first 3 d after the OM application. With one exception, > 50% of the rebuilt macroaggregates were formed within the first 7 d after rewetting and addition of OM. However, the amount of organic C within the new macroaggregates was ≈ 2‐ to 3‐fold higher than in the original soil. The process of aggregate formation was still proceeding after 7 d of incubation, however at a lower rate. Contents of organic C within macroaggregates were decreased markedly after 28 d of incubation in the OM₁ and OM₂ treatments, suggesting that the microbial biomass (bacteria and fungi) used organic C within the newly built macroaggregates. Overall, the results confirmed for all treatments that macroaggregate formation is a rapid process and highly connected with the amount of OM added and microbial activity. However, the time of maximum aggregation after C addition depends on the soil and substrate investigated. Moreover, the results suggest that the primary macroaggregates, formed within the first 7 d, are still unstable and oversaturated with OM and therefore act as C source for microbial decomposition processes.     

川西平原土壤微生物生物量碳氮磷含量特征及其影响因素分析

本文通过区域调查采样和统计分析,探讨了川西平原土壤微生物生物量碳（MBC）、土壤微生物生物量氮（MBN）和土壤微生物生物量磷（MBP）含量特征及其对气候、海拔、母质和土地利用等因素的响应,揭示了其关键影响因素,以期为川西平原地区土壤质量管理提供参考。结果表明,不同土壤类型的MBC、MBN和MBP含量表现为冲积土显著高于水稻土、潮土和黄壤（P<0.05）,潮土MBC/MBN显著高于水稻土。气候和海拔的影响为：MBC、MBN和MBP含量随着≥ 0℃积温、≥ 10℃积温、年均温和年均降水量的增加呈指数减少,而随干燥度和海拔增加呈线性增加。不同成土母质中,MBC、MBN和MBP含量均为灰色冲积物显著高于老冲积物。不同土地利用方式下,三者含量为草地显著高于水田和旱地,水田、旱地和林地差异不显著。皮尔森相关分析和冗余分析表明,MBC和MBN均与≥ 0℃积温、年均温呈极显著负相关（P<0.01）,与海拔呈极显著正相关关系,MBP与母质呈现极显著负相关关系。逐步回归分析表明,MBC主要受年均温、干燥度、年均降水量和母质的影响;MBN主要受海拔、干燥度和年均降水量的综合影响;MBP主要受母质、年均温、≥ 10℃积温和年均降水量的调控。因此,川西平原土壤MBC、MBN、MBP能灵敏地反映不同采样点气候的变化,可为该区气候变化下土壤碳、氮、磷的响应预测提供参考。     

Effect of land degradation on soil microbial biomass in a hilly area of south Sumatra,Indonesia

We investigated the impact of land-use changes on the soil biomass at several soil sites in Indonesia under different types of land-use (primary forest, secondary forest, coffee plantation, traditional orchard, and deforested area), located within a small geographical area with similar parent material and climatic conditions. Various parameters of soil microbial biomass (biomass C, biomass N, content of anthrone-reactive carbohydrate carbon, and soil ergosterol content) were examined. Our results suggested that the removal of the natural plant cover did not cause any appreciable decrease in the amount of microbial biomass; on the contrary it led to a short-time increase in the amount of microbial biomass which may be due to the availability of readily decomposable dead roots and higher sensitivity to the decomposition of residual litter in recently deforested soils. However, the amount of microbial biomass tended to decrease in proportion to the duration of the land history in coffee plantation soils. This may be ascribed to the effect of the loss of available substrates associated with soil erosion in the long term. Lower ergosterol contents in recently deforested areas reflected a reduction in the amount of fungal biomass which may be due to the destruction of the hyphal network by the slash and burn practice. On the other hand, the higher soil ergosterol content at the sites under bush regrowth indicated that microbial biomass was able to recover rapidly with the occurrence of a new plant cover.     

Long‐term effects on soil microbial properties of heavy metals from industrial exhaust deposition

Soil samples were taken at 0—10 cm and 10—20 cm depth from 7 clay‐marsh sites used as grassland close to Nordenham in the north of Lower Saxony, Germany. The sites had been contaminated by deposition of heavy metals from industrial exhausts, the level of contamination varying according to their distances from a lead factory. The soils were analyzed to assess the depth‐specific effects of NH₄NO₃ extractable and total amounts of Zn, Pb, and Cu on basal respiration, adenylates, ergosterol, and biomass C estimated by fumigation extraction (FE) and substrate‐induced respiration (SIR). Most of the chemical and biological properties studied decreased with depth, but depth‐specific differences in the relationships between these properties rarely occurred. The biomass C/soil organic C ratio was at a relatively high level, but most consistently reflected pollution as a decrease with increasing heavy metal load, independently of the method used for biomass C estimation. However, the SIR estimates were on average 44 % lower than those of FE, mainly due to pH effects. The metabolic quotient SIR‐qCO₂ increased with increasing NH₄NO₃ extractable and total heavy metal contents, but also with decreasing pH, whereas the FE‐qCO₂ remained unaffected by heavy metals and pH. The ATP/FE‐biomass C ratio was on average 8.2 μmol g^—1 and negatively affected by soil pH, but also by total Zn, NH₄NO₃ extractable Zn and Cu. The ergosterol/FE‐biomass C ratio was on average 0.29 %, i.e. at a very low level, and increased with increasing heavy metal content. This indicates a change in the community structure towards fungi.     

Soil microorganisms and the growth of Lupinus albus on a high metal soil in the presence of EDTA

A pot experiment was designed with the objective of determining whether the presence of ethylenediaminetetraacetic acid (EDTA) and the resulting mobilization of heavy metals have any affect on: (i) soil microorganisms, (ii) growth of L. albus, and (iii) microbial colonization of roots. There was no effect of the different treatments on the contents of soil microbial biomass C and microbial biomass N. Increasing addition of EDTA to soil led to proportionate increases in extractable C and N, being roughly equivalent to the added amount. Increasing EDTA addition to soil led also to a proportionate increase in mobile heavy metals. Plant height, total amount of shoot and root C were not affected by EDTA addition. Fungal ergosterol in the lupine roots showed a 5- to 8-fold increase in the 0.1 and 0.2% EDTA treatments in comparison with the control. In contrast, EDTA addition did not affect fungal glucosamine or bacterial muramic acid concentrations in the lupine roots. Increasing EDTA addition to soil led also to a proportionate increase in the metal concentrations in the lupine shoots. The concentrations of heavy metals in the lupine shoots and in the NH₄NO₃ soil extracts were all highly significantly correlated.     

Gel formation mechanism and gel properties controlled by Ca2+ in chia seed mucilage and model substances

Polygalacturonic acid (PGA) is considered as a model substance for mucilage to study mucilage–soil interactions, assuming that the gel formation mechanism of mucilage is comparable to the one of PGA. However, some studies question the accepted hypothesis, which states that, like for PGA, this mechanism relies on cross‐links between uronic acid and calcium for mucilage. The aim of this study was therefore to understand the influence of the abundance and degree of esterification of uronic acids and the influence of calcium on the gel formation mechanism in mucilage as compared to model substances. The mucilage used was from chia seeds, as it is easily available in great quantity and has gel properties shared by root mucilage. Results reported here demonstrate that, while the gel formation mechanism of PGA relied on specific cross‐links with calcium and led to heterogeneous gels, low‐methoxy pectin (LMP) formed homogeneous calcium gels also characterized by nonspecific ionic interactions with calcium. On the contrary, despite similar uronic acid content to LMP, chia seed mucilage was mostly governed by weak electrostatic interactions between entangled polymer chains, which conferred the gel poor water retention. Addition of calcium reduced repulsion and molecular expansion, resulting in a reduction of the water content in chia seed mucilage. Finally, the discrepancies between PGA, LMP and chia seed mucilage discredit the use of PGA as model for chia seed mucilage. Comparison with root mucilage is still needed. This study offers the keys for further mechanistic understanding on the influence of mucilage on soil properties.     

Coffee response to nitrogen and soil water content during the early growth stage

Coffee (Coffea arabica L.) is the major crop grown in highland and mountainous areas of Colombia, where the most common yield‐limiting factors are nitrogen (N) and soil water content. Since the eventual success of a plantation is determined by the performance of coffee plants during critical early stages, our goal was to better understand plant response to these limiting factors. Four different levels of N and soil water content were evaluated under greenhouse conditions for their effect on growth, water use efficiency, fertilizer‐N recovery efficiency (FNRE), and stem lignin and cellulose of coffee seedlings. The interaction between N and moisture did not produce a significant response in coffee growth. However, by increasing N, both water use efficiency and ¹³C content were enhanced, while growth, recovery of urea by plants, and stem strength were decreased. Water stress due to low soil water content increased water use efficiency, ¹³C content, and root to shoot ratio, but decreased shoot growth. These results demonstrate the effects of N supply and water balance and highlight the excessive amount of N typically applied to coffee seedlings as well as the importance of the acclimation process of young plants to changes in soil N and water. All of these are important considerations in improving management strategies to reduce environmental impact while sustaining optimal productivity.     

Appropriate nitrogen supply could improve soil microbial and chemical characteristics with Sophora davidii seedlings cultivated in water stress conditions

Abstract

A greenhouse experiment was conducted to investigate the changes of soil microbial activities and chemical properties under different water and nitrogen (N) supply conditions. A completely randomized design was subjected to three water regimes (80%, 40% and 20% water field capacity (FC)) and three N supply regimes (control, N0: 0 mg N kg^?1 soil; low N supply, Nl: 92 mg N kg^?1 soil; and high N supply, Nh: 184 mg N kg^?1 soil) by potting with 2-month-old Sophora davidii seedlings. Water stress decreased the content of soil organic carbon (C), available N and phosphorus (P), the ratio of C/N, the ratio of C/P, as well as activities of soil invertase, urease and alkaline phosphatase, but not reduced microbial biomass C, N and P contents. Soil microbial and chemical characteristics also exhibited strong responses to N supply, and these responses were inconsistent among N supply levels. The contents of soil organic C and available P showed stronger positive responses to Nl than to Nh, while the available N content increased with increasing N supply. Additionally, Nl rather than the other two N treatments led to increased microbial biomass N and invertase activity under 20% FC treatment, even though the invertase activity increased in Nh treatment under 40% FC and 80% FC treatments. Nl treatment also increased the C/P ratio and alkaline phosphatase activity. These results suggest that water and N co-limited nutrient mineralization and microbial activity, and that these characteristics responded positively to Nl. Therefore, appropriate or low N supply is recommended to increase soil quality restrained by water stress, thereby facilitating S. davidii seedling establishment under water deficit conditions.     

水氮耦合对盐碱地柳枝稷生物质产量、品质及水肥利用的影响

在宁夏银北盐碱地区,以'Cave-in-rock'柳枝稷为材料,采用随机区组试验设计,研究了水氮耦合对柳枝稷生物质产量、品质及水肥利用效率的影响,利用模糊相似优先比法分析了不同水氮处理下柳枝稷产量、品质及水肥利用效率的变化特征.结果表明,在盐碱地条件下,充足的灌水和适量的氮肥投入是柳枝稷高产和水肥高效利用的保证.在相同...     

Effects of land-use and management practices on soil ergosterol content in andosols

Ergosterol is a fungus-specific chemical component which has been extensively used to assess the fungal biomass and activity in soils. In the present study, we investigated the soil ergosterol content in a group of Japanese soils differing in management. In addition, the relationships among the soil ergosterol content (SEC), microbial biomass C (MBC), and microbial biomass N (MBN) were also tested in different seasons. Our results showed that the SEC was lower in intensively cropped soils than in pasture and forest soils. In general, no consistent trend was observed for SEC of pasture soils within the same sampling period. However, SEC tended to decrease in most of the soils in autumn, which may be ascribed to the effect of seasonal changes on fungal biomass. No strong evidence could be obtained for the effect of different fertilizers on SEC, which can be due to different management variables and interspecific variations between fungal communities at different soil sites. Our results did not suggest the presence of a noticeable correlation between SEC and MBC or MBN. The ratio of SEC in total biomass C was lower (0.20 and 0.11% in spring and autumn, respectively) compared with the values reported.     

An efficient method for the collection of root mucilage from different plant species—A case study on the effect of mucilage on soil water repellency

Root mucilage may play a prominent role in understanding root water uptake and, thus, there is revived interest in studying the function of root mucilage. However, mucilage research is hampered by the tedious procedures of mucilage collection. We developed a mucilage separator which utilizes low centrifugal forces (570 rpm) to separate the mucilage from seminal roots without the need of handling individual seeds or removing the germinated seeds from the tray/mesh to a centrifuge tube. For the different plant species, between 1 and 3.7 mL tray^?1 of hydrated mucilage could be produced, with 6 trays being handled successively within 45 min. For Triticum aestivum, which showed a dry matter content of 0.5%, this was equivalent to 98.6 mg mucilage dry matter. The lowest total production was found for Zea mays with just 34 mg dry matter. The amounts of mucilage produced normalized to root tip agree well with literature data. The mucilage obtained by the new method was used to measure its effect on repellency of soil as this property directly relates to the phenomenon of lower rhizosphere soil water content during rewetting. It could be shown that repellency of the rhizosphere is affected by the quantity as well as by species‐dependent quality of mucilage in the rhizosphere. Among the species tested (Lupinus albus, Vicia faba, Zea mays, Triticum aestivum), the largest differences were observed between the two legumes. For Zea mays seminal root mucilage obtained with the new system was compared to mucilage of air born brace roots. The differences between these two mucilages, representing different root orders, indicate clearly that there is still a need for methods which enable the investigation of roots from older plants.     

鸡粪与化肥不同配比对杨树苗根际土壤酶和微生物量碳、氮变化的影响

为比较鸡粪有机肥与化肥以不同比例配施对杨树苗根际土壤环境的作用效果,通过盆栽试验,研究了N100（尿素提供100%的氮）、 M10N90（鸡粪和尿素分别提供10%和90%的氮)、 M30N70（鸡粪和尿素分别提供30%和70%的氮）和M50N50（鸡粪和尿素各提供50%的氮）等处理对一年生杨树苗根际土壤酶活性、 微生物量碳（SMBC）和微生物量氮（SMBN）的影响。结果表明, 同N100处理相比,M30N70处理的杨树苗根际土壤脲酶、 碱性磷酸酶、 过氧化氢酶和蔗糖酶活性分别提高14.41%~26.49%、 13.87%~27.93%、 12.07%~27.08%和29.17%~46.43%; 根际SMBC、 SMBN含量分别提高63.00%~76.62%、 40.01%~90.38%。在杨树苗的年生长过程中,根际土壤脲酶、 碱性磷酸酶、 过氧化氢酶和蔗糖酶活性均在施肥后120 d 最高; 而根际SMBC和SMBN含量分别在施肥后90 d和30 d达到最大值。随着杨树苗的生长发育,SMBN含量逐渐降低,而在落叶期有一定的回升。与M30N70处理相比,M10N90和M50N50处理对杨树苗根际土壤酶和微生物量碳、 氮的影响较小。综合分析认为,鸡粪有机肥与化肥以3∶7比例配施对杨树苗根际区域生态环境的改善效果最佳。     

黑河流域保护性耕作对农田土壤微生物量碳、氮和有机质的调控

为了探讨黑河流域保护性耕作对土壤生产力的影响,设计20cm留茬(NS20),20cm留茬压倒(NPS20),40cm留茬(NS40),40cm留茬压倒(NPS40)和传统耕作(CT)5个处理,研究了黑河流域保护性耕作对农田土壤有机质、土壤微生物量C、土壤微生物量N以及作物产量和水分利用效率的影响。结果表明,保护性耕作农田0—20cm土层土壤有机质、土壤微生物量C和N的含量均高于传统耕作,且其在剖面中的变化趋势基本一致,即随土层深度增加下降;土壤微生物量N有明显的"表聚现象";相关分析表明土壤有机质和土壤微生物量C之间显著正相关(r=0.85,p0.05),与土壤微生物量N之间无明显的相关关系(r=0.47,p0.05);保护性耕作提高了春小麦的产量,NPS20和NPS40增产效果最好,较CT分别增产53.08%和46.59%,与CT之间差异达到极显著水平;保护性耕作提高了春小麦的水分利用效率(WUE),NPS20,NS40,NPS40,NS20分别较CT的WUE提高了58.02%,43.40%,47.27%,23.78%。     

Microcalorimetric evaluation of soil microbiological properties under plant residues and dogmatic water gradients in Red soil

A study of 6 months duration was carried out to investigate the effect of water regimes and organic amendments on the soil microbial biomass and microbial population under Red soil collected from Hunan Providence, China. The soil microbial biomass and population were measured with traditional methods and results obtained by conventional methods, corroborated with microcalorimetry. The incorporation of rice (Oryza sativa L.) straw (RS) and green manure (GM), especially at high rates, enhanced the soil microbial activities. We observed that the use of GM exhibited more significant stimulating effects on microbial activities than RS. Similarly, water regimes, 25% (W1) and 200% (W2) of water holding capacity, also had significant effects on microbial activities. Comparing the effects of water levels, we noticed that W2 had a significant negative influence on soil microbial biomass and population. To compare the results of conventional methods and to check the sensitivity of microcalorimetry, the thermodynamic parameters, microbial growth rate constant (k), total heat evolution (Q), peak height (P_max) and peak time (t_max) were calculated. Highest P_max, k and Q were observed in GM treatments at water regime W1, while highest t_max values were recorded in CK (control) and RS treatments at W2. The microcalorimetric parameters, P_max, k and Q were positively correlated, whereas t_max negatively linked with microbial biomass and population at p < 0.01. Our results suggest that microcalorimetry successfully verified the results obtained from customary methods and microcalorimetric parameters P_max, t_max, Q and k proved that they are highly sensitive to microbial properties and could be used as indices of microbial community shifts and activities in soil ecosystems.     

Using metabolic tracer techniques to assess the impact of tillage and straw management on microbial carbon use efficiency in soil

Tillage practices and straw management can affect soil microbial activities with consequences for soil organic carbon (C) dynamics. Microorganisms metabolize soil organic C and in doing so gain energy and building blocks for biosynthesis, and release CO₂ to the atmosphere. Insight into the response of microbial metabolic processes and C use efficiency (CUE; microbial C produced per substrate C utilized) to management practices may therefore help to predict long term changes in soil C stocks. In this study, we assessed the effects of reduced (RT) and conventional tillage (CT) on the microbial central C metabolic network, using soil samples from a 12-year-old field experiment in an Irish winter wheat cropping system. Straw was removed from half of the RT and CT plots after harvest or incorporated into the soil in the other half, resulting in four treatment combinations. We added 1-¹³C and 2,3-¹³C pyruvate and 1-¹³C and U-¹³C glucose as metabolic tracer isotopomers to composite soil samples taken at two depths (0–15 cm and 15–30 cm) from each of the treatments and used the rate of position-specific respired ¹³CO₂ to parameterize a metabolic model. Model outcomes were then used to calculate CUE of the microbial community. Whereas the composite samples differed in CUE, the changes were small, with values ranging between 0.757 and 0.783 across treatments and soil depth. Increases in CUE were associated with a reduced tricarboxylic acid cycle and reductive pentose phosphate pathway activity and increased consumption of metabolic intermediates for biosynthesis. Our results suggest that RT and straw incorporation do not substantially affect CUE.