Physiological water strategy of Artemisia ordosica around soil threshold of drought

The mechanism for plants around the soil threshold of drought (close to the soil wilting water content) is a problem that needs to be further explored. In this paper, Artemisia ordosica, which grows in the Tengri Desert, was selected to analyze the changes in the plant water potentials in the soil-plant-atmosphere continuum (SPAC), the water contents in the roots, shoots and leaves of A. ordosica, and the indices in enzymatic and non-enzymatic systems. Based on the statistics, we discussed the water physiology mechanism around the soil drought threshold. The results show that, around the soil drought threshold, besides absorbing and transporting water, the roots could serve as temporary water reservoirs that enable A. ordosica to continue to transport the SPAC water and survive severe drought. As drought becomes more severe, the activity of superoxide dismutase (SOD) and catalase (CAT) increases and they have significant correlations with the tissue water content. The activity of peroxidase (POD) decreases and it has no significant correlation with the tissue water content. During daytime, when temperature is high, the soluble sugar does not participate in the osmotic adjustment but eliminate the active oxygen free radicals. Thus, around the soil threshold of drought, A. ordosica maintains a physiological water metabolism by harmonizing water itself and eliminate the active oxygen and the free radicals by the joint efforts of enzymatic and nonenzymatic systems.     

EFFECTS OF AM FUNGI AND WATER STRESS ON DROUGHT RESISTANCE OF ARTEMISIA ORDOSICA IN DIFFERENT SOILS

 利用两种不同土壤研究了水分胁迫和接种AM真菌(摩西球囊霉(Glomus mosseae)和油蒿(Artemisia ordosica)根际土著AM真菌)对毛乌素沙地重要演替物种油蒿生长和抗旱性的影响。结果表明, 两种土壤中水分胁迫没有显著影响油蒿的植株形态和含水量, 但严重抑制了菌根侵染率。水分胁迫促使油蒿提高叶片保水能力, 抑制N、P在地上部的分配。在胁迫前期SOD活性较高, 而POD活性在后期较高。同一水分条件下接种AM真菌显著提高了AM真菌侵染率, 土壤中孢子数显著增多, 提高了植株分枝数并促进侧根发育, 显著提高根冠比和植株保水能力, 加强了根系对全磷、全氮的吸收。接种AM真菌的植株可溶性糖和丙二醛含量较低, 可溶性蛋白含量无显著变化, SOD和POD活性提高, 油蒿抗旱性加强。水分胁迫下在不同土壤中接种不同AM真菌对油蒿的促进效应差异较大, 接种土著AM真菌的效果优于摩西球囊霉单一接种。干旱导致菌根侵染率下降是宿主植物吸水能力下降的原因之一, 在植物生长前期接种AM真菌可以增强植物抵抗生长中后期环境干旱的能力。     

干旱胁迫下AM真菌对油蒿叶片保护系统的影响

基利用盆栽试验在正常水分和干旱胁迫条件下研究了灭菌土接种AM真菌摩西球囊霉(Glomus mosseae)和土著AM真菌对油(蒿Artemisia ordosica)生长及叶片保护系统的影响。结果表明,干旱胁迫显著抑制了土著AM真菌对油蒿的侵染,但对G.mosseae的侵染影响较小。正常水分和干旱胁迫条件下,接种AM真菌显著增加了油蒿生物量和干重以及根系含磷量;提高了叶绿素、可溶性糖、可溶性蛋白含量并降低了脯氨酸和丙二醛含量;显著增强了过氧化氢酶(CAT)和过氧化物酶(POD)活性,增强了油蒿对干旱的防御能力。     

植物水分传输过程中的调控机制研究进展

农田土壤水分的转化利用与调控是以土壤-植物-大气连续体(SPAC)为基础,以植物为核心,其中水分在植物体内的传输与调控研究一直是国际学术研究的前沿性热点课题。本文概述了植物水分传输的驱动力和传输途径,重点从植物的气孔调节、水容调节、渗透调节、水孔蛋白调节、贮水调节、气穴和栓塞调节等方面综述了植物水分传输过程中的调控机制研究进展。通过对植物存在优化调控水分平衡的潜在能力的研究,不仅可充实SPAC系统水分传输理论,而且有助于明确植物对环境的适应机制和高效用水的潜力及其节水调控的效应,对指导干旱半干旱地区农业生产提供理论依据。     

不同降水条件下两种荒漠植物的水分利用策略

自然降水是干旱、半干旱地区荒漠植物重要的水分来源。为了说明自然降水量的变化对干旱、半干旱地区荒漠植物水分利用策略的影响, 研究了两种常见荒漠植物油蒿(Artemisia ordosica)和白刺(Nitraria tangutorum)在3个不同自然降水地区(内蒙古的杭锦旗和磴口县及甘肃的民勤县)的水分来源、水分利用效率及植物的抗逆能力的变化。测定了不同地区的植物茎水、各潜在水源(降水、地下水和土壤水)的δD和δ¹⁸O值, 并利用IsoSource模型分析了这两种植物在不同地区对这些潜在水源的选择性利用情况; 同时测定了叶片的δ¹³C和游离脯氨酸浓度。结果表明: 在年降水量最高的杭锦旗, 这两种植物对浅层土壤水的利用比例最高, 其中油蒿主要利用0-50 cm土层中的水源; 在年降水量相对较低的磴口和民勤, 植物利用的主要水源为深层土壤水和地下水。随着年降水量的增加, 这两种植物的水分利用效率逐渐降低。白刺的脯氨酸浓度大于油蒿, 与水分利用效率无关, 但油蒿的水分利用效率和脯氨酸浓度成正比。研究表明, 荒漠植物能通过改变其水分利用策略和其他生理特性适应自然降水量的变化, 但不同植物种采用的策略可能有所不同。     

Saikosaponin accumulation and antioxidative protection in drought-stressed Bupleurum chinense DC. plants

Dried root of Bupleurum spp. is one of the most popular ingredients in many oriental medicinal preparations. Potted Bupleurum chinense DC. seedlings were subjected to progressive drought stress by withholding irrigation followed by a rewatering phase. The changes in antioxidant system, hydrogen peroxide (H₂O₂), superoxide radicals (O₂^?), and malondialdehyde (MDA) contents as well as saikosaponin a (SSa) and saikosaponin d (SSd) content in B. chinense roots were investigated. Additionally, the antioxidant activity of the roots extract was evaluated. The results showed that B. chinense root appeared highly resistant to water deficit. Both SSa and SSd content increased with the progressive water deficit, however, decreased under severe drought conditions or after water recovery. Moderate drought treatment resulted in 83% increase in SSa content and 22% increase in SSd content compared to the well-hydrated treatment. And increased SSa and SSd content during drought were accompanied by enhanced O₂^? content and superoxide dismutase (SOD), catalase (CAT), ascorbate peroxidase (APX) activity until severe drought stress. Notably, in vitra antioxidant tests demonstrated that the lipid peroxidation inhibition capacity was positively correlated with the content of SSa and SSd, particularly significant at p = 0.05 with SSd content. These results suggest that B. chinense roots exhibit effective antioxidative protection mechanism to withstand drought stress. And it could be speculated that drought-induced SSa and SSd accumulation in B. chinense roots may be stimulated via active oxygen species, and consequently involve in mitigating the oxidative damage due to its high anti-lipid peroxidation capacity.     

农林复合系统水分生态特征的模拟研究

根据验证后的太行山低山丘陵区苹果 -小麦复合系统及单作小麦系统 SPAC水分运移模型 ,模拟计算小麦拔节 -腊熟期间复合系统和单作系统蒸腾、根系吸水和土壤水分的时空变化值 ,继而分析复合系统水分生态特征 ,旨在为该地区发展果农复合模式提供一定的理论依据 ,并以期进一步丰富复合农林业水分生态理论研究内容。结果表明 :(1)对比单作小麦系统 ,复合系统具有明显的小麦蒸腾需水及耗水降低效应 ,其值分别可达 18.0 %和 16 .8% ;实际区域面积上 ,复合系统中作物和果树蒸腾耗水量的比值为 1.7,说明小麦耗水是复合系统耗水的主要特征方式。 (2 )分布林带区和作物区的果树吸水量分别约占系统内果树吸水总量的 4 7.6 %和 5 2 .4 % ,说明果树所消耗的土壤水分至少有 5 0 %以上来自作物区内 ;在作物与果树根系的交错区内 ,作物吸水量与果树吸水量的比值达 82 .0 :1,因此 ,小麦在与果树根系吸水的直接竞争过程中 ,将处绝对主导的地位。 (3)小麦拔节-腊熟期间各时段内 ,复合系统中 0～ 80 cm土层贮水量随带距的变化均大致呈抛物线状分布 ,对比单作小麦系统 ,复合系统土壤水分总体平均效应 2 .7%。     

Generation of Active Oxygen Free Radicals and Its Injury to Microsome Membranes in Wheat Leaves Under Drought Stress

The microsome membrane of wheat ( Triticum aestivum L. ) leaves under different time of drought stress was purified to determine the active oxygen free radicals generated and their severity of injury to the membrane. During the drought stress the rate of O2- and H2O2 generation increased for a certain time and then decreased. The concentration of MDA increased continuously. The activity of SOD decreased gradually. The concentration of —SH group and MDA increased from the beginning until the 36th hour under drought stress, thereafter MDA content continued to increase while the —SH content decreased. Several physiological indexes related to membrane injury, such as root electroconductivity, leaf water potential and water content were all affected accordingly.     

Nitrate uptake ability by maize roots during and after drought stress

The effects of different intensities and durations of soil drought and re-watering on the nitrate uptake ability of maize roots were studied. Plants were grown in split-root containers with one part of the root system subjected to different intensities and durations of soil drought and re-watering while the other part of the root system was continuously watered to 23% (w/w) soil water content (70% water capacity). Experiments were performed in split-root containers to maintain a high growth rate, thus ensuring high nutrient demand of the shoot irrespective of the soil water regime. To avoid limitation of nitrate uptake by transport processes in the dry soil, and to ensure a uniform ¹⁴N/¹⁵N ratio at the root surface, ¹⁵N was applied to the roots by placing them into an aerated nutrient solution with 0.5 mM Ca(¹⁵NO₃)₂. Shoot elongation and biomass were only slightly affected by drought in one root compartment when the soil in the other root compartment was kept wet. Therefore, the growth-related nutrient demand of the shoot remained at a high level. At moderate levels of soil drought (10% w/w water content) the ability of the roots for N-uptake was not affected even after 10 d of drought. N-uptake ability was reduced to about 20% of the well-watered control only when the soil water content was decreased to 5%. Total soluble sugar content of the roots increased with increasing soil drought, indicating that low N-uptake ability of roots subjected to severe soil drought was not caused by low assimilate supply from the shoot. Nitrate uptake ability of roots maintained in very dry soil (5% soil water content w/w) even for a prolonged period of 8 d, recovered within 3 d following re-watering. Root growth increased one day after re-watering. A short-term experiment with excised roots formerly subjected to severe soil drought showed that nitrate uptake ability recovered in old and young root segments after 2 d of re-watering. Obviously, the increase in N-uptake ability after re-watering was caused not only by new root growth but also by recovery of the uptake ability of formerly stressed roots.     

不同配置模式下油蒿和柠条对降雨变化的生理生态响应及调节机制

研究了沙坡头地区固沙植物油蒿和柠条在1 m×1 m和2 m×2 m配置模式下的群落学特征,以及在自然和控制降雨条件下的水分关系、气体交换及其调节机制.结果表明： 随着土壤含水量的变化,油蒿和柠条的生理耗水和生长特性不同,油蒿的气孔导度、水势、蒸腾速率、光合速率和快速生长均呈现明显的波动性,并且对土壤水分的消耗量高于柠条;柠条的上述指标变化相对低缓而恒定,光合生长速率较低,多年累积生物量却很高.两种植物对水分胁迫的响应过程和调节机制不同,油蒿主要为保水机制,柠条主要为节水机制.在极端干旱年份,柠条的节水、抗逆特性更强.固沙植物种的选择应既利于防风固沙和节约用水,又利于固沙林的稳定.     

干旱胁迫下桑树叶片中超氧阴离子的变化规律

该研究以6个四川桑树品种为对象,对供试桑树品种进行连续的人工干旱处理,利用生理方法明确保护酶系(SOD)的变化规律,并采用硝基四氮唑蓝(NBT)还原试验进行超氧阴离子O_2~-·组织化学的可视化检测。结果表明:随着干旱胁迫的加剧,SOD的含量明显高于对照组,其中SOD活性呈先升后降的变化趋势,而超氧阴离子一直呈加重的趋势。6个桑树品种的SOD含量从正常到中度胁迫都呈现不同程度的上升趋势,上升趋势明显的品种是九龙拐和湖桑32,上幅分别为62.7%和60.1%,其次是云桑1号和荷叶白一号上幅,分别为52.2%、58.2%,油桑和充桑的上幅为50.8%和47.4%,说明SOD正发挥着积极的作用。在中度胁迫之后SOD的含量与活性氧的水平表现出相反的趋势,原因可能是随着干旱胁迫的加剧,O_2~-·达到最大值且超出一定域值,从而导致SOD活性的降低。通过电镜观察发现,超氧阴离子由叶脉向周围扩散,同时面积逐渐增大。其中,抗旱性强的品种湖桑32叶片表面茸毛比较长,且基部膨大,端部有明显的刺状突起,而抗旱性弱的品种荷叶白1号叶片表面茸毛比较细短且端部的刺状突起不明显,因此推测叶片表面茸毛的特性与抗旱性存在一定关系。     

干旱胁迫下小麦叶片微粒体活性氧自由基的产生及其对膜的伤害

从不同干旱胁迫时间下的小麦(Triticum aestivum L.)叶片中提取微粒体膜,测定活性氧自由基变化和膜伤害程度。结果表明,随着干旱胁迫时间的延长发生以下变化:(1)O_2~-和H_2O_2的产生速率在一定时间内逐渐加快然后下降;(2)膜脂过氧化产物丙二醛(MDA)的含量逐渐增加;(3)微粒体膜上的SOD活性逐渐下降;(4)—SH基的含量在一定时间内逐渐增多以后下降;(5)与膜伤害程度有关的生理指标如根系电导率、叶片水势及其含水量等均有变化。     

活性氧、自由基与植物的衰老

介绍近 1 0年来有关活性氧、自由基的产生 ,对植物的伤害及植物对活性氧、自由基清除的研究进展。     

春小麦对骤旱的响应特征及其阈值分析

与缓慢发展的干旱过程不同,骤旱具有发生速度快,短期内可致害的特点。目前,关于作物骤旱致害的临界阈值及其调控机制尚不清楚。以春小麦为供试作物,通过桶栽试验,模拟研究骤旱过程中小麦受旱致害的过程特征及其控制因素。结果发现,发生骤旱时土壤含水量下降呈先快后慢的变化趋势,叶片水分和叶水势则呈先慢后快的指数变化趋势。叶片光合生理指标对土壤水分的下降存在明显的阈值响应,且不同生理指标的阈值并不完全相同,其中净光合速率与表征叶片光合能力的指标（最大羧化速率）对土壤有效含水量的响应阈值为0.4,气孔导度和蒸腾速率对土壤有效含水量的响应阈值分别为0.5和0.4。而小麦光合生理指标对叶片水分和叶水势的阈值响应并不明显。同时依据各生理指标相关和通径分析结果得出,骤旱发生时引起小麦叶片净光合速率快速降低的主导因子为非气孔因素,而并不是以往作物受旱研究中的气孔因素。本研究结果有望丰富干旱影响认知,并可为科学应对干旱提供依据。     

不同干旱土壤条件下杨树的耗水规律及水分利用效率研究

 在适宜土壤水分（70%θf）,中度干旱（55%θf）和严重干旱（40%θf）3种土壤水分条件下研究杨树（Populus simonii）的耗水特性和水分利用特征。结果表明,随着土壤含水量的下降,杨树叶水势、相对含水量(RWC)、生长速率、光合速率及单叶水分利用效率（WUE）显著下降;在适宜水分和中度干旱条件下,杨树的快速生长和干物质迅速积累时期主要集中在5～6月,严重干旱下快速生长时期和干物质积累主要集中在5月;杨树总耗水量和总生物量的大小顺序为：适宜水分＞中度干旱＞严重干旱;WUE则表现出中度干旱下最高,严重干旱下最低;杨树在适宜水分下的日、旬、月耗水量明显高于中度干旱和严重干旱处理;杨树在适宜水分、中度干旱和严重干旱条件下的最高耗水月分别在6～7月,最高旬耗水量分别在7月中旬、上旬和6下旬;在中度水分亏缺和严重水分亏缺下的最高耗水日出现的时间比适宜水分下的最高耗水日提前1～2个月以上。一天中的最大耗水高峰随着杨树生育期和土壤含水量的不同而有明显差异。研究结果表明,杨树不具备耐旱植物的特征,因此在黄土高原缺水地区不适宜大面积栽植,只能用于水分条件较好的立地条件下造林。     

短期干旱对水稻叶水势、光合作用及干物质分配的影响

采用盆栽水分试验,研究了不同生育期短期干旱处理对水稻叶水势、光合作用和干物质分配的影响．结果表明,干旱胁迫后,水稻叶水势低于对照,午后叶水势回升缓慢。凌晨叶水势随土壤含水量的降低而降低,表现为阈值反应。叶片净光合速率与凌晨叶水势密切相关,低于凌晨叶水势临界值,水稻叶片净光合速率急剧下降在水稻抽穗期和灌浆期叶片净光合速率显著下降的凌晨叶水势临界值为-1.04和-1．13MPa,对应的土壤含水量阈值分别为饱和含水量的61.0％和50．9％,土壤水势分别为-0．133和-0．240MPa干旱胁迫下单叶净光合速率的日变化规律表现为：胁迫较轻时,单叶净光合速率在正午附近出现低谷;胁迫严重时,净光合速率全天低于对照,且不及对照的一半。短期干旱后,水稻叶、根、穗的分配指数均降低,茎鞘的分配指数升高。本研究可为水稻节水灌溉管理和水分限制下水稻的生长模拟提供生理基础和理论依据。     

海滨滨麦叶片和根对不同厚度沙埋的生理响应差异分析

以烟台海岸抗风沙植物滨麦为研究材料,通过对不同厚度沙埋下其叶片和根部抗氧化酶活力(超氧化物歧化酶(SOD)、过氧化物酶(POD)、过氧化氢酶(CAT))、丙二醛(MDA)含量和渗透调节物含量变化的分析,探讨了叶片和根部对沙埋生理响应的差异。试验按滨麦成株株高(约40 cm)对其进行了轻度沙埋(在株高1/4处)、中度沙埋(2/4处)和重度沙埋(3/4处)。在沙埋第6天,分别测定了不同厚度沙埋处理下,植株各段叶片和根抗氧化酶活力、MDA和渗透调节物含量。结果表明,轻度和中度沙埋均加速植株生长。与对照相比,经轻度、中度沙埋处理6 d,叶片平均MDA含量增加,在重度沙埋下降低。不同厚度沙埋6 d,叶片平均SOD活力和脯氨酸含量增加,而CAT活力、可溶性糖和可溶性蛋白质含量下降。但不同厚度沙埋均使沙上叶片MDA、脯氨酸、可溶性蛋白质含量和SOD和CAT活力增加,尤其是叶片顶部增加最为明显,使沙下叶片MDA、可溶性糖、可溶性蛋白质含量和CAT活力下降,导致同株沙上和沙下叶片MDA、脯氨酸、可溶性糖、可溶性蛋白质含量和SOD和CAT活力差异显著(P0.05)。与叶片相比,根中MDA、可溶性蛋白质含量和SOD和CAT活力较低,而POD活力和可溶性糖含量较高并与叶片差异显著(P0.05)。不同厚度沙埋6 d,滨麦根中MDA和可溶性蛋白质含量变化较小,可溶性糖含量和CAT、POD、SOD活力略有降低。研究表明,滨麦根和叶片对不同厚度沙埋的生理响应不同。沙埋直接作用于叶片并诱发叶内氧自由基积累,但叶片通过快速激活的抗氧化酶保护系统(CAT、SOD)维持氧自由基代谢平衡,以及渗透调节物(脯氨酸、可溶性糖)的积累维护细胞水分代谢平衡,并满足能量的需求和快速生长。但在不同厚度沙埋下,由于根系不受沙埋直接影响而生理变化较小,并且还维持较低的膜脂过氧化水平,这可能是根能维持正常的吸水输水功能并在沙埋处理过程中和沙埋后地上叶片快速生长摆脱沙埋的重要物质基础。     

土壤-根系统水分再分配:土壤-植物-大气连续体中的一个小通路

吸收和传导水分一直被视为植物根系最主要的功能之一,而人们对根系在某些情况下还可以向土壤释放水分的事实及其对植物生长和生态系统功能的影响了解得还很不充分,尽管这样的证据由来已久。土壤-根系统水分再分配(Hydraulic redistribution, HR)是近20年间被发现和证实的,指水分从土壤中较湿的部分经由植物的根系传导而运动到土壤中较干的部分,通常发生在蒸腾减弱的夜间,可以沿水势梯度下降的方向而在不同土层间向上向下或侧向运动。HR研究揭示了土壤-植物-大气连续体中有时会存在土壤-根-土壤的水流小通路,细化了土壤-根系统中水分储存和运输的时空动态和机制。土壤水分状况的连续监测、根木质部液流测量、稳定性同位素技术的使用构成了HR实验研究的三大手段。当土壤中深层水分充足的时候,HR可以提高根系吸收和传导水分的效率,有利于植物充分利用资源,延长了浅层土壤的水分可利用期,有利于维持植物组织的生理活性和水流传导;旱季后降水来临的时候,HR可以将一部分降水转移到深层土壤,增加了可利用性水分的总量。对于干旱半干旱的沙地和草原、季节性干旱的森林等类型,HR过程可能对生态系统水分循环产生重要影响。有必要在国内针对这些生态系统展开深入的实验研究,同时探索将HR过程适当结合到生态系统模型和水文模型中,从而更准确地研究和预测群落内植物水分关系和生态系统水分动态。此外,结合农林设计、植被恢复、生态需水量估算和农业节水等方面进行的HR研究也值得深入探索。     

水分胁迫对春播玉米苗期生长及其生理生化特性的影响

采用盆栽试验,以‘蠡玉18'玉米单交种为供试材料,设置充分供水(CK)、轻度水分胁迫(LS)、中度水分胁迫(MS)和重度水分胁迫(SS)4个水分处理水平,研究了水分胁迫对春播玉米苗期保护酶活性和生长的影响,以探讨土壤水分胁迫对玉米苗期生长发育及其生理过程的影响机制.结果表明:(1)随着水分胁迫程度的加剧,玉米幼苗的生物量显著下降,根冠比、根系活力和脯氨酸含量增加,且水分胁迫对玉米幼苗地上部生物量的抑制作用更大;可溶性蛋白含量差异不明显,MDA含量波动变化.(2)随着水分胁迫时间的延长,根冠比、根系活力和植株脯氨酸含量先升高后降低,可溶性蛋白含量呈先下降后升高的趋势;玉米幼苗叶片和根系MDA积累波动变化,而叶片MDA含量始终高于根系.(3)在水分胁迫初期,玉米叶片中CAT活性较SOD、POD响应更敏感;玉米苗期根系在中度水分胁迫下主要依赖CAT来降低氧化危害,而在重度水分胁迫下前期主要依赖CAT、后期通过CAT和POD的共同作用来降低氧化伤害;水分胁迫条件下,叶片和根系POD同步降低氧化伤害,而SOD和CAT在叶片和根系间存在互补作用.研究表明,在不同程度的水分胁迫条件下,玉米幼苗的生长受到一定程度的抑制,但其能够通过调节自身的保护酶活性和渗透调节物质含量来减轻干旱伤害,维持植株的正常生理代谢功能.     

Effects of Cadmium on Enzymatic and Non-Enzymatic Antioxidative Defences of Rice (Oryza Sativa L.)

The effects of 60-d cadmium (Cd) exposure on enzymatic and non-enzymatic antioxidative system of Oryza sativa L. seedlings at tillering stage were studied using soil culture experiment. Research findings showed that chlorophyll content of Oryza sativa L. declined with the increase in soil metal concentration. Cd pollution induced the antioxidant stress by inducing O₂ ^?1 and H₂O₂, which increased in plants; at the same time, MDA as the final product of peroxidation of membrane lipids, accumulated in plant. The antioxidant enzyme system was initiated under the Cd exposure, i.e. almost all the activities of superoxide dismutase (SOD), peroxidase, catalase, glutathione peroxidase, and ascorbate peroxidase were elevated both in leaves and roots. The non-protein thiols including phytochelatins and glutathione to scavenge toxic free radicals caused by Cd stress was also studied. The contents of phytochelatins and glutathione were about 3.12–6.65-fold and 3.27–10.73-fold in leaves, against control; and the corresponding values were about 3.53–9.37-fold and 1.41–5.11-fold in roots, accordingly.