Soil microbial community responses to altered lignin biosynthesis in Populus tremuloides vary among three distinct soils

The development and use of transgenic plants has steadily increased, but there are still little data about the responses of soil microorganisms to these genetic modifications. We utilized a greenhouse trial approach to evaluate the effects of altered stem lignin in trembling aspen (Populus tremuloides) on soil microbial communities in three soils which differed in their chemical and physical properties; they included a sandy loam (CO-Colorado), a silt loam (KS-Kansas), and a clay loam (TX-Texas). Three transgenic aspen lines were developed from a natural clone common to the Great Lakes region of North America. The concentrations of stem lignin concentrations were reduced by 35% (Line 23), 40% (Line 141) and 50% (Line 72). Line 72 and Line 141 also had a 40 and 20% increase in syringyl-type stem lignin, respectively. Indirectly, these modifications resulted in increased (5–13%) and decreased (−5 to −57%) levels of root production across the lines and soil types. Responses of the soil microbial communities were investigated using: phospholipid fatty acids (PLFA), neutral lipid fatty acids (NLFA), and 3) extracellular enzyme assays. PLFA analyses indicated that there were large differences in microbial community composition between the three soils. Similarly, there were large differences in total NLFA between soils, with the KS soils having the highest amount and CO the lowest. Enzyme activities did not differ between soils, except for cellubiohydrolase, which was highest in CO soil. Across all three soils, responses to the four genetic lines were not consistent. Interactions between soil type and genetic line make it difficult to assess the potential ecological impacts of transgenic aspen on soil microbial communities and their associated functions. Given these interactions, field trials with transgenic aspen should encompass the wide range of soils targeted for commercial planting in order to determine their effect(s) on the resident soil microbial community. Responsible Editor: Barbara Wick     

The influence of soil type and altered lignin biosynthesis on the growth and above and belowground biomass allocation of Populus tremuloides

Plants influence soil carbon (C) formation through the quality and quantity of C released to soil. Soil type, in turn can modify a plant’s influence on soil through effects on plant production, tissue quality and regulation of soil C decomposition and stabilization. Wild-type aspen and three transgenic aspen lines expressing reduced stem lignin concentrations and/or increased syringyl (S) to guaiacyl (G) ratio lignin were grown in greenhouse mesocosms containing a sandy loam, a silt loam, or a clay loam soil for 6 months in order to examine the effects of altered lignin biosynthesis and soil type on biomass partitioning (above vs. belowground) and soil C processes. Results indicated that soil type significantly affected plant performance. Aspen grown in soils with high sand/low clay content accumulated the most total biomass, while aspen grown in soils with high clay content accumulated the least total biomass. These reductions in growth combined with specific soil characteristics led to differences among soil types in soil C formation. Transformed aspen expressing high syringyl/guaiacyl (S/G) lignin accumulated less total plant C and subsequently accumulated less aspen derived C in soil. Reduced lignin content alone in aspen did not affect plant growth or soil C formation. There were significant soil type × genetic line interactions indicating that growth and soil C formation for transgenic and wild type aspen lines varied among the different soil types. Given these interactions, future investigation needs to include long-term field studies across a range of soil types before transgenic aspen are widely planted.     

Factors affecting fall down rates of dead aspen (Populus tremuloides) biomass following severe drought in west‐central Canada

Increases in mortality of trembling aspen (Populus tremuloides Michx.) have been recorded across large areas of western North America following recent periods of exceptionally severe drought. The resultant increase in standing, dead tree biomass represents a significant potential source of carbon emissions to the atmosphere, but the timing of emissions is partially driven by dead‐wood dynamics which include the fall down and breakage of dead aspen stems. The rate at which dead trees fall to the ground also strongly influences the period over which forest dieback episodes can be detected by aerial surveys or satellite remote sensing observations. Over a 12‐year period (2000–2012), we monitored the annual status of 1010 aspen trees that died during and following a severe regional drought within 25 study areas across west‐central Canada. Observations of stem fall down and breakage (snapping) were used to estimate woody biomass transfer from standing to downed dead wood as a function of years since tree death. For the region as a whole, we estimated that >80% of standing dead aspen biomass had fallen after 10 years. Overall, the rate of fall down was minimal during the year following stem death, but thereafter fall rates followed a negative exponential equation with k = 0.20 per year. However, there was high between‐site variation in the rate of fall down (k = 0.08–0.37 per year). The analysis showed that fall down rates were positively correlated with stand age, site windiness, and the incidence of decay fungi (Phellinus tremulae (Bond.) Bond. and Boris.) and wood‐boring insects. These factors are thus likely to influence the rate of carbon emissions from dead trees following periods of climate‐related forest die‐off episodes.     

The response of sensitive and tolerant clones of Populus tremuloides to dynamic ozone exposure under controlled environmental conditions

Both sensitive and tolerant clones of aspen ( Populus tremuloides ) were exposed to ozone using four different exposure regimes under controlled environmental conditions. Based on data on ambient ozone from 10 cities in the USA, three treatments of 4-wk exposure to the same SUM06 (an accumulation of hourly O₃ concentrations greater than 0.06 ml l⁻¹) were constructed. The regimes allowed us to investigate: (a) the importance of long (3 wk, treatment 1) versus short (1 wk, treatment 2) duration of regimes with high peaks; (b) the effect of treatments with variable peak occurrence (treatments 1 and 2) versus uniform peak occurrence (treatment 3) during the exposure period. Nonfumigated control plants were maintained at ozone concentrations <10 nl l⁻¹. Bifacial black necrosis, a typical symptom of ozone injury on aspen leaves, occurred on both clones after 2 wk exposure. Up to 60% of the leaves on the sensitive clone were injured, with an average of 6% of total leaf area injured. In the tolerant clone only 10% of the leaves were injured, with less than 1% of the total leaf area symptomatic. The severity of injury was consistently greatest in treatment 2, followed by treatments 1 and 3, respectively. The interval between peak exposures was less important than the occurrence of peaks versus a stable maximum concentration. Premature leaf abscission occurred in the sensitive clone. Measures of gas exchange demonstrated reduced photosynthesis under ozone fumigation, but exposure regime was not a significant factor. Concentrations of two antioxidants, ascorbic acid and glutathione, were almost always greater in the resistant than in the sensitive clone, but the differences were not statistically significant. The levels of these antioxidants in aspen leaves did not change with ozone fumigation or leaf age.     

Competition- and resource-mediated tradeoffs between growth and defensive chemistry in trembling aspen (Populus tremuloides)

Costs of defense are thought to maintain genetic variations in the expression of defense within plant populations. As with many plant species, aspen exhibits considerable variation in allocation to secondary metabolites. This study examined the independent and interactive effects of genotype, soil fertility and belowground competition on defensive chemistry and growth in trembling aspen (Populus tremuloides). Four aspen genotypes were grown with high and low soil fertility, and with and without root competition. Physiological, morphological and allocational determinants of growth were measured to identify growth-defense tradeoffs. Nutrient limitation and competition decreased growth, leaf mass ratio, leaf nitrogen concentration and photosynthesis, and increased root : shoot ratio and leaf condensed tannin concentrations. The competition treatment also resulted in increased leaf phenolic glycoside (PG) concentrations. Aspen growth was negatively correlated with PG concentrations under low fertility with competition. The relationship between growth and its major determinants was also negatively related to foliar condensed tannins expressed as a proportion of tree mass, indicating an additional indirect cost of allocation to secondary metabolites.     

Physiological and biomass partitioning shifts to water stress under distinct soil types in Populus deltoides saplings

土壤类型是影响植物分布和生产力的重要环境因素,但有关土壤环境异质性对植物抗逆性效应的研究非常缺乏。本研究以美洲黑杨 (Populus deltoids)为对象,以3种典型土壤类型(红壤、黄壤和黄棕壤)为栽培基质,在控制实验条件下,经过三个月的干旱胁迫(25%田间持水量)处理,测定了不同处理条件下美洲黑杨的气体交换速率、抗氧化能力、氮代谢特征、生物量积累与分配特征。研究结果表明,在红壤和黄壤条件下,与对照(75%田间持水量)相比,干旱胁迫显著降低了美洲黑杨各器官的生物量、光合速率、叶片氮同化酶的活性,显著增 加了叶片中过氧化氢、丙二醛和无机氮的含量。在黄棕壤条件下,干旱对美洲黑杨总生物量、光合速率、氮同化酶以及质膜完整性的负面 影响较小,这与其维持较高的超氧化物歧化酶,过氧化物酶和过氧化氢酶的活性相关,也与其生物量分配模式(如提高根冠比)密切相关。 由此可见,生长在黄棕壤条件下的美洲黑杨表现出较强的抗旱能力,这可能与其土壤母质中较高的土壤养分和良好的通气状况相关。因此,就土壤类型而言,与红壤和黄壤相比,黄棕壤提供的土壤环境条件有利于美洲黑杨的抗逆表现和栽培利用。     

Stem wood properties of Populus tremuloides, Betula papyrifera and Acer saccharum saplings after 3 years of treatments to elevated carbon dioxide and ozone

The aim of this study was to examine the effects of elevated carbon dioxide [CO₂] and ozone [O₃] and their interaction on wood chemistry and anatomy of five clones of 3‐year‐old trembling aspen (Populus tremuloides Michx.). Wood chemistry was studied also on paper birch (Betula papyrifera Marsh.) and sugar maple (Acer saccharum Marsh.) seedling‐origin saplings of the same age. Material for the study was collected from the Aspen Free‐Air CO₂ Enrichment (FACE) experiment in Rhinelander, WI, USA, where the saplings had been exposed to four treatments: control (C; ambient CO₂, ambient O₃), elevated CO₂ (560 ppm during daylight hours), elevated O₃ (1.5 × ambient during daylight hours) and their combination (CO₂+O₃) for three growing seasons (1998–2000). Wood chemistry responses to the elevated CO₂ and O₃ treatments differed between species. Aspen was most responsive, while maple was the least responsive of the three tree species. Aspen genotype affected the responses of wood chemistry and, to some extent, wood structure to the treatments. The lignin concentration increased under elevated O₃ in four clones of aspen and in birch. However, elevated CO₂ ameliorated the effect. In two aspen clones, nitrogen in wood samples decreased under combined exposure to CO₂ and O₃. Soluble sugar concentration in one aspen clone and starch concentration in two clones were increased by elevated CO₂. In aspen wood, α‐cellulose concentration changed under elevated CO₂, decreasing under ambient O₃ and slightly increasing under elevated O₃. Hemicellulose concentration in birch was decreased by elevated CO₂ and increased by elevated O₃. In aspen, elevated O₃ induced statistically significant reductions in distance from the pith to the bark and vessel lumen diameter, as well as increased wall thickness and wall percentage, and in one clone, decreased fibre lumen diameter. Our results show that juvenile wood properties of broadleaves, depending on species and genotype, were altered by atmospheric gas concentrations predicted for the year 2050 and that CO₂ ameliorates some adverse effects of elevated O₃ on wood chemistry.     

Populus tremuloides photosynthesis and crown architecture in response to elevated CO2 and soil N availability

We tested the hypothesis that elevated CO₂ would stimulate proportionally higher photosynthesis in the lower crown of Populus trees due to less N retranslocation, compared to tree crowns in ambient CO₂. Such a response could increase belowground C allocation, particularly in trees with an indeterminate growth pattern such as Populus tremuloides. Rooted cuttings of P. tremuloides were grown in ambient and twice ambient (elevated) CO₂ and in low and high soil N availability (89 ± 7 and 333 ± 16 ng N g⁻¹ day⁻¹ net mineralization, respectively) for 95 days using open-top chambers and open-bottom root boxes. Elevated CO₂ resulted in significantly higher maximum leaf photosynthesis (A _max) at both soil N levels. A _max was higher at high N than at low N soil in elevated, but not ambient CO₂. Photosynthetic N use efficiency was higher at elevated than ambient CO₂ in both soil types. Elevated CO₂ resulted in proportionally higher whole leaf A in the lower three-quarters to one-half of the crown for both soil types. At elevated CO₂ and high N availability, lower crown leaves had significantly lower ratios of carboxylation capacity to electron transport capacity (V _cmax/J _max) than at ambient CO₂ and/or low N availability. From the top to the bottom of the tree crowns, V _cmax/J _max increased in ambient CO₂, but it decreased in elevated CO₂ indicating a greater relative investment of N into light harvesting for the lower crown. Only the mid-crown leaves at both N levels exhibited photosynthetic down regulation to elevated CO₂. Stem biomass segments (consisting of three nodes and internodes) were compared to the total A _leaf for each segment. This analysis indicated that increased A _leaf at elevated CO₂ did not result in a proportional increase in local stem segment mass, suggesting that C allocation to sinks other than the local stem segment increased disproportionally. Since C allocated to roots in young Populus trees is primarily assimilated by leaves in the lower crown, the results of this study suggest a mechanism by which C allocation to roots in young trees may increase in elevated CO₂. Received: 12 August 1996 / Accepted: 12 November 1996     

Plant leaf traits,height and biomass partitioning in typical ephemerals under different levels of snow cover thickness in an alpine meadow

Aims In the cold life zones, snow cover is a comprehensive environmental factor that directly influences soil temperature, soil water content, light and nutrient availability. Plants in these zones develop a series of unique mechanisms involving phenological characteristics, reproductive strategies, physiology and morphology to adapt to environmental changes. This paper is focused on the responses of plant leaf traits, height and biomass partitioning to variations in snow cover thickness, in order to better understand the responses of plant functional traits and specific adaptation strategies under global climate change scenarios. Methods Three transects were established along a gradient of snow cover in an alpine meadow of Mt. Kaka, in the eastern Qinghai-Xizang Plateau. Primula purdomii, Pedicularis kansuensis and Ranunculus tanguticus, which are three widely distributed and dominant ephemerals, were sampled and studied, particularly at their blooming stages. Plant height, specific leaf area (SLA) and biomass partitioning were measured accordingly. Important findings The values of SLA in Pedicularis kansuensis and R. tanguticus were relatively greater under better soil conditions; it was smaller in Primula purdomii with thick snow cover. The relationship between aboveground biomass and belowground biomass in Primula purdomii was allometric at sites with both thick and thin snow cover. No significant relationships were found between aboveground biomass and belowground biomass in Pedicularis kansuensis and R. tanguticus at some individual sites. However, when samples of the three species were pooled, the relationships between aboveground biomass and belowground biomass were allometric at all sites, which did not support isometric scaling hypothesis. In addition, on sites with either thick or thin snow cover, aboveground biomass had greater rate of accumulation than belowground biomass; whereas on sites with medium snow cover, the rate of biomass accumulation was greater for belowground component than aboveground component. Functional traits and biomass variables were better correlated in Primula purdomii and Pedicularis kansuensis than in R. tanguticus.     

不同雪被厚度下典型高山草地早春植物叶片性状、株高及生物量分配的研究

在高寒生命带, 雪被作为重要的综合环境因子, 影响着植物的生理生态特征、种群动态及群落演替等过程, 进而作用于生态系统的功能与服务。通过在青藏高原东缘高寒草甸设置厚雪、中雪和浅雪3个地段, 选取早春开花的常见种紫罗兰报春(Primula purdomii)、甘肃马先蒿(Pedicularis kansuensis)、高原毛茛(Ranunculus tanguticus), 研究了三种植物株高、叶片性状和生物量分配随雪被厚度的变化规律, 以及三者之间的关系。结果表明: 甘肃马先蒿和高原毛茛在生境状况较好的地段比叶面积相对较高, 紫罗兰报春由于具有根状茎和肉质根, 在厚雪地段比叶面积相对较小; 关于单个物种的地上-地下生物量的关系, 紫罗兰报春在厚雪和浅雪地段为完全一致的异速生长关系, 而甘肃马先蒿和高原毛茛在个别地段并无显著相关关系。总体而言, 三种植物整体样本的地上-地下生物量在不同雪被地段均为异速生长关系, 不支持等速生长假说, 并且地上生物量能够很好地解释地下生物量的变异; 功能性状和生物量指标间的相关性, 在紫罗兰报春和高原毛茛表现较好, 而在甘肃马先蒿表现较弱。植物对环境变化的适应具有一定的规律, 同时也表现出物种特异性。今后的研究中, 应注重构建植物适应环境变化的功能性状谱, 以更好地理解全球变化背景下植物功能性状的响应及其适应策略。     

Field‐grown transgenic switchgrass (Panicum virgatum L.) with altered lignin does not affect soil chemistry,microbiology, and carbon storage potential

Cell wall recalcitrance poses a major challenge on cellulosic biofuel production from feedstocks such as switchgrass (Panicum virgatum L.). As lignin is a known contributor of recalcitrance, transgenic switchgrass plants with altered lignin have been produced by downregulation of caffeic acid O‐methyltransferase (COMT). Field trials of COMT‐downregulated plants previously demonstrated improved ethanol conversion with no adverse agronomic effects. However, the rhizosphere impacts of altering lignin in plants are unknown. We hypothesized that changing plant lignin composition may affect residue degradation in soils, ultimately altering soil processes. The objective of this study was to evaluate effects of two independent lines of COMT‐downregulated switchgrass plants on soils in terms of chemistry, microbiology, and carbon cycling when grown in the field. Over the first two years of establishment, we observed no significant differences between transgenic and control plants in terms of soil pH or the total concentrations of 19 elements. An analysis of soil bacterial communities via high‐throughput 16S rRNA gene amplicon sequencing revealed no effects of transgenic plants on bacterial diversity, richness, or community composition. We also did not observe a change in the capacity for soil carbon storage: There was no significant effect on soil respiration or soil organic matter. After five years of establishment, δ¹³C of plant roots, leaves, and soils was measured and an isotopic mixing model used to estimate that 11.2 to 14.5% of soil carbon originated from switchgrass. Switchgrass‐contributed carbon was not significantly different between transgenic and control plants. Overall, our results indicate that over the short term (two and five years), lignin modification in switchgrass through manipulation of COMT expression does not have an adverse effect on soils in terms of total elemental composition, bacterial community structure and diversity, and capacity for carbon storage.     

Photosynthesis,respiration, and carbon allocation of two cool-season perennial grasses in response to surface soil drying

The study was conducted to investigate carbon metabolic responses to surface soil drying for cool-season grasses. Kentucky bluegrass (Poa pratensis L.) and tall fescue (Festuca arundinaceae Schreb.) were grown in a greenhouse in split tubes consisting of two sections. Plants were subjected to three soil moisture regimes: (1) well-watered control; (2) drying of upper 20-cm soil (upper drying); and (3) drying of whole 40-cm soil profile (full drying). Upper drying for 30 d had no dramatic effects on leaf water potential (Ψ_leaf) and canopy photosynthetic rate (P_n) in either grass species compared to the well-watered control, but it reduced canopy respiration rate (R_canopy) and root respiration rate in the top 20 cm of soil (R_top). For both species in the lower 20 cm of wet soil, root respiration rates (R_bottom) were similar to the control levels, and carbon allocation to roots increased with the upper soil drying, particularly for tall fescue. The proportion of roots decreased in the 0-20 cm drying soil, but increased in the lower 20 cm wet soil for both grass species; the increase was greater for tall fescue. The Ψ_leaf, P_n, R_canopy, R_top, R_bottom, and carbon allocation to roots in both soil layers were all significantly higher for upper dried plants than for fully dried plants of both grass species. The reductions in R_canopy and R_top in surface drying soil and increases in root respiration and carbon allocation to roots in lower wet soil could help these grasses cope with surface-soil drought stress. This revised version was published online in June 2006 with corrections to the Cover Date.     

Productivity, leaf traits and carbon isotope discrimination in 29 Populus deltoides x P. nigra clones

Here we tested whether some leaf traits could be used as predictors for productivity in a range of Populus deltoides x P. nigra clones. These traits were assessed in 3-yr-old rooted cuttings from 29 clones growing in an open field trial, in a five randomized complete block design, under optimal irrigation. Variables were assigned to four groups describing productivity (above-ground biomass, total leaf area), leaf growth (total number of leaves increment rate), leaf structure (area of the largest leaf, specific leaf area, carbon and nitrogen contents), and carbon isotope discrimination in the leaves (Delta). High-yielding clones displayed larger total leaf area and individual leaf area, while no correlation could be detected between productivity and either leaf structure or Delta. By contrast, Delta was negatively correlated with number of leaves increment rate and leaf N content. Our study shows that there is a potential to improve water-use efficiency in poplar without necessarily reducing the overall productivity.     

Increased microbial growth,biomass, and turnover drive soil organic carbon accumulation at higher plant diversity

Species‐rich plant communities have been shown to be more productive and to exhibit increased long‐term soil organic carbon (SOC) storage. Soil microorganisms are central to the conversion of plant organic matter into SOC, yet the relationship between plant diversity, soil microbial growth, turnover as well as carbon use efficiency (CUE) and SOC accumulation is unknown. As heterotrophic soil microbes are primarily carbon limited, it is important to understand how they respond to increased plant‐derived carbon inputs at higher plant species richness (PSR). We used the long‐term grassland biodiversity experiment in Jena, Germany, to examine how microbial physiology responds to changes in plant diversity and how this affects SOC content. The Jena Experiment considers different numbers of species (1–60), functional groups (1–4) as well as functional identity (small herbs, tall herbs, grasses, and legumes). We found that PSR accelerated microbial growth and turnover and increased microbial biomass and necromass. PSR also accelerated microbial respiration, but this effect was less strong than for microbial growth. In contrast, PSR did not affect microbial CUE or biomass‐specific respiration. Structural equation models revealed that PSR had direct positive effects on root biomass, and thereby on microbial growth and microbial biomass carbon. Finally, PSR increased SOC content via its positive influence on microbial biomass carbon. We suggest that PSR favors faster rates of microbial growth and turnover, likely due to greater plant productivity, resulting in higher amounts of microbial biomass and necromass that translate into the observed increase in SOC. We thus identify the microbial mechanism linking species‐rich plant communities to a carbon cycle process of importance to Earth's climate system.     

Overexpression of Populus trichocarpa CYP85A3 promotes growth and biomass production in transgenic trees

Brassinosteroids (BRs) are essential hormones that play crucial roles in plant growth, reproduction and response to abiotic and biotic stress. In Arabidopsis, AtCYP85A2 works as a bifunctional cytochrome P450 monooxygenase to catalyse the conversion of castasterone to brassinolide, a final rate‐limiting step in the BR‐biosynthetic pathway. Here, we report the functional characterizations of PtCYP85A3, one of the three AtCYP85A2 homologous genes from Populus trichocarpa. PtCYP85A3 shares the highest similarity with AtCYP85A2 and can rescue the retarded‐growth phenotype of the Arabidopsis cyp85a2‐2 and tomato d^x mutants. Constitutive expression of PtCYP85A3, driven by the cauliflower mosaic virus 35S promoter, increased the endogenous BR levels and significantly promoted the growth and biomass production in both transgenic tomato and poplar. Compared to the wild type, plant height, shoot fresh weight and fruit yield increased 50%, 56% and 43%, respectively, in transgenic tomato plants. Similarly, plant height and stem diameter increased 15% and 25%, respectively, in transgenic poplar plants. Further study revealed that overexpression of PtCYP85A3 enhanced xylem formation without affecting the composition of cellulose and lignin, as well as the cell wall thickness in transgenic poplar. Our finding suggests that PtCYP85A3 could be used as a potential candidate gene for engineering fast‐growing trees with improved wood production.     

荒漠土壤微生物量碳、氮变化对降水的响应

以腾格里沙漠东南缘的典型荒漠植被为研究对象,通过遮雨棚和滴灌系统设置5个降水梯度,即极端干旱处理、中度干旱处理、对照、增水处理I和增水处理II,研究了荒漠土壤微生物量碳(MBC)、氮(MBN)和微生物碳氮比(MBC/MBN)对季节、降水和土壤深度的响应规律,以期为极端降水事件影响干旱荒漠区土壤微生物量碳、氮及其循环规律的深入研究提供科学依据。结果表明:(1)MBC、MBN和MBC/MBN对降水处理的响应存在差异,三者的变化范围为:230.14—272.87 mg/kg,13.82—17.58 mg/kg,19.78—36.06。其中,降水处理对MBC、MBN的影响显著,对MBC/MBN的影响不显著,在极端干旱处理下,MBC、MBN均显著高于其他降水处理;(2)两年间的MBC、MBN和MBC/MBN差异显著,2017年较2016年MBC、MNB显著减少,MBC/MBN显著增加;(3)MBC、MBN和MBC/MBN变化均表现季节性差异,变化范围分别为:153.31—337.09 mg/kg,7.89—22.29 mg/kg,14.82—46.04,其中MBC、MBN为春季最高、秋季最低,M...