Relationship between root elongation rate and diameter and duration of growth of lateral roots of maize

The objective of this work was to describe the relationship between elongation rate and diameter of maize roots and to estimate the length and growth duration of lateral roots of maize. Diameters and elongation rates of roots were measuredin situ on plants grown 5 weeks in small rhizotrons under greenhouse conditions. At the end of the experimental period the roots were harvested and diameters of axile and lateral roots were measured. The frequency distribution of diameters of harvested roots was bimodal with a minimum at 0.6 mm; 97% of axile roots were larger than this value and 98% of the lateral roots were smaller. Root elongation per day increased as diameter increased but the slope of the relationship with lateral roots was about 2.5 times that with axile roots when separate linear regressions were fitted to the two populations. The length of lateral roots found on axillary roots between the base and about 30 cm from the apex was approximately 2.2 cm. All of the data was consistent with the hypothesis that the lateral roots grew for about 2.5 days and then ceased growing. The axillary roots continued to grow throughout the experimental period at a rate of about 3 cm day⁻¹. Contribution from the Department of Agronomy, New York State College of Agriculture and Life Sciences, Cornell University, Ithaca, NY 14853. Agronomy paper No. 1661. This research is part of the program of the Center for Root-Soil Research.     

Applications of functional classification methods for tree fine root biomass estimation: Advancements and synthesis

The morphology of fine root branching of woody plants is highly variable in their forms and functions. In the past two decades, researchers have increasingly recognized that the root-diameter-based method, using an arbitrary size of root diameter, failed to precisely characterize the physiological and ecological processes involved in finest roots. The number of publications using root-order-based approaches has increased regardless the fact that root trait-measurements based on root order are time-consuming and labor-intensive. A new approach—root functional classification method—was proposed and had been applied in the literature. The functional classification of fine roots separates roots of < 2 mm to absorptive and transport pools, making it more feasible for studies on root biomass and turnover. This new concept redefines fine root guild and has great potentials for future studies. Our literature review of the topic indicates that less is known about the inter-specific differences in estimates of biomass of absorptive and/or transport roots, with a large variation of absorptive roots on global scale. In addition, our review emphasizes the importance in: a) precision estimating of the absorptive biomass of fine roots, and b) proper definition of the range of the transport roots within and among forest ecosystems. Finally, after compare the strengths and weaknesses of the functional classification method, we propose several specific suggestions to improve the applications of this approach.     

The Developmental Anatomy of the Seminal Root of Wheat

Wheat embryo usually gives rise to five seminal roots in matured caryopsls, although, the sixth root might develop in some cases. The first one is known as the primary root. Primary root emerged early, and its primodium was distinctly originated from the proembryo and could be gradually identified as three layers of initials. Lateral seminal roots emerged later from the embryonic axis in pairs, and originated from the surrouding cells of the procambium. Differentiation of lateral roots was much more vigorous than that of the first seminal root (primary root), and, its mother cells of metaxylem vessel appeared soon, Lateral seminal roots usually had more metaxylem vessels. In short, only the first root is the primary root, the lateral seminal roots are adventitious in nature, since their structures are similar to those of other adventitious roots.     

Using a fine root extraction device to quantify small diameter corn roots (≥0.025 mm) in field soils

The study of fine roots growing under field conditions is limited by the techniques currently available for separating these roots from soil. This study had two objectives: to measure the total root length of field grown corn (Zea mays L.) by root diameter class, and to develop an inexpensive and efficient root washing device that would effectively capture all of the roots in a field soil sample. An inexpensive Fine Root Extraction Device (FRED) was constructed from readily available materials and was successful at extracting all roots, including very fine diameter roots (0.025 mm), from field soil samples. Greater than 99.7% of marked roots introduced to the FRED were recaptured by the device. Soil samples from three depths, and on three dates, from field grown corn were placed in the FRED. We found that more than 56% of total root length occurred in roots whose diameters were smaller than 0.175 mm, and more than 35% of root length occurred in roots smaller than 0.125 mm in diameter. Corn roots of the diameters described here have not been reported in field soils prior to this study. Root researchers who fail to measure these very fine roots will significantly underestimate root length density. Widespread use of the FRED should improve our understanding of root distribution in field soils.     

玉米初生根和不同层次次生根的直径和大量元素含量的比较分析

对大田条件下玉米品种'掖单13'(Zea mays 'Yedan 13')和'丹玉13'(Z. mays 'Danyu 13')初生根(包括初生胚根和次生胚根)和不同层次次生根的直径及其中的N、P、K、Ca和Mg含量进行了比较分析.分析结果表明,'掖单13'和'丹玉13'初生根直径总体上小于次生根;从第1层至第7层次生根,基本上为发生时间越晚,根直径越大;而2个品种第7层至第9层次生根基部直径的变化趋势有一定的差异.'掖单13'和'丹玉13'初生根和不同层次次生根中N、P、K、Ca和Mg含量差异显著(P<0.05);'掖单13'初生根中的N、P、K、Ca和Mg含量分别为27.36、 3.75、 14.34、 10.19和5.94 g·kg~(-1), 不同层次次生根中的N、 P、 K、 Ca和Mg含量分别为4.31～8.99、 0.91～1.29、4.07～15.27、2.35～5.42和1.52～4.06 g·kg~(-1);'丹玉13' 初生根中的N、P、K、Ca和Mg含量分别为28.65、4.65、14.54、10.33和6.04 g·kg~(-1),不同层次次生根中的N、P、K、Ca和Mg含量分别为7.41～12.70、0.84～1.27、1.81～24.69、2.46～3.73和1.12～2.73 g·kg~(-1);2个品种初生根中N、P、Ca和Mg含量均最高,K含量也显著高于第1层至第7层次生根;总体上看,不同层次次生根中的N、Ca和Mg含量随发生时间的先后呈逐渐降低的趋势,P和K含量随发生时间的先后呈逐渐增加的趋势,但2个品种间有一定的差异.研究结果显示,次生根根直径的增大与其对地上部分的固持能力有关.     

小麦种子根的发育解剖

小麦胚胎发育过程中通常形成5条幼根(少数可形成6条),这些根统称为种子根,中间最先发生的为初生根.初生根的原基在胚胎发育的早期就在胚轴的一侧发生,原基细胞由不规则到规则排列。侧生种子根的原基在胚胎发育后期才出现,通常成对发生,并且是由胚轴上的节(盾片节和胚芽鞘节)维管束外方的细胞形成。侧生种子根的发育明显较初生根的快,分化能力也较强,后生木质部导管母细胞出现早,数目较多.因此,小麦胚胎发育过程中从胚轴上形成的这些侧生的种子根,形态上,仍应看作是一些不定根,其结构特征与后来形成须根系的不定根的比较近似。     

青杨人工林根系生物量、表面积和根长密度变化

在植物生长季节,采用钻取土芯法对秦岭北坡50年生青杨人工林根径≤2 mm和2～5 mm根系的生物量、表面积和根长密度进行测定.结果表明：在青杨人工林根系(＜5 mm)中,根径≤2 mm根系占总生物量的77.8%,2～5 mm根系仅占22.2%;根径≤2 mm根系表面积和根长密度占根系总量的97%以上,而根径2～5 mm根系不足3%.随着土层的加深,根径≤2 mm根系生物量、表面积和根长密度数量减少,根径2～5 mm根系生物量、表面积和根长密度最小值均分布在20～30 cm土层.≤2 mm根系生物量、表面积和根长密度与土壤有机质、有效氮呈极显著相关,而根径2～5 mm根系的相关性不显著.     

川中丘陵区3个树种的细根形态和功能异质性分析

以川中丘陵区柏木低效林林窗改造初期种植的银木和香椿细根为研究对象,以未改造的柏木纯林为对照,采用LI-8100土壤碳通量测量系统测定银木、香椿和柏木1~5级细根的原位呼吸速率,并探讨细根形态结构和养分元素浓度与细根呼吸的相关关系,以揭示细根结构与功能异质性。结果表明:银木、香椿和柏木细根的直径、根长、组织碳浓度均随着根序级别的增加而增加,而它们细根的比根长、组织氮浓度和比根呼吸速率均随着根序的增加而降低,树种、根序级及其交互作用对3个树种细根形态、养分浓度和比根呼吸均有显著或极显著影响。回归分析显示,3个树种比根呼吸速率均随细根直径、比根长、N浓度变化呈现出系统性的变化,三者分别能解释64.7%、87.6%和、67.6%的比根呼吸变异。可见,细根在形态和功能上存在明显的异质性,且细根的形态特征、组织化学含量和生理功能之间存在着紧密的联系,为理解植物根系结构与功能变异提供了依据。     

Changes in the risk of fine-root mortality with age: a case study in peach, Prunus persica (Rosaceae)

Previous studies suggest that younger roots are more vulnerable to mortality than older roots. We analyzed minirhizotron data using a mixed-age, proportional hazards regression approach to determine whether the risk of mortality (or "hazard") was higher for younger roots than for older roots in a West Virginia peach orchard. While root age apparently had a strong effect on the hazard when considered alone, this effect was largely due to different rates of mortality among roots of different orders, diameters, and depths. Roots with dependent laterals (higher order roots) had a lower hazard than first-order roots in 1996 and 1997. Greater root diameter was also associated with a decreased hazard in both 1996 and 1997. In both years, there was a significant decrease in the hazard with depth. When considered alone, age appeared to be a strong predictor of risk: a 1-d increase in initial root age was associated with a 1.26-2.62% decrease in the hazard. However, when diameter, order, and depth were incorporated into the model, the effect of root age disappeared or was greatly reduced. Baseline hazard function plots revealed that the timing of high-risk periods was generally related to seasonal factors rather than individual root age.     

高水位条件下池杉根系的生态适应机制和膝根的呼吸特性

  池杉(Taxodium ascendens)属于典型的耐水树种, 掌握其根系对淹水环境的生态适应机制对于研究林木耐水机理十分重要。通过对江苏省里下河低湿地17年生池杉在高水位(6～10月淹水, 全年平均地下水位-5 cm)、中水位(8～9月淹水, 全年平均地下水位-18 cm)和低水位(常年不淹水, 全年平均地下水位-41 cm)条件下的根系进行调查, 结果表明, 池杉在高水位条件下形成细长的气生根, 气生根依附于树干北侧或潜伏于树干外表皮内侧和纵裂的树皮缝隙中; 中水位池杉能形成直径(7.9±2.2) cm、高(7.7±2.7) cm的膝根, 每株立木拥有膝根数(5.8±1.7)个; 低水位池杉也能形成膝根, 但个体小、数量少。林木地下和地上生物量均呈现出明显的高水位<中水位<低水位的趋势, 但是地下/地上生物量的比值却呈相反趋势, 表明池杉耐水性虽然很强, 长期处于较高水位时生长会明显受抑, 尤其是地上生物量生长受抑更显著。高、中和低水位池杉的地径/胸径之比分别是2.66±0.11、2.08±0.10和1.75±0.08, 说明水位较高的环境能促进树干基部的相对粗生长。长期淹水导致地下根的容重降低, 但是气生根和膝根的容重却明显大于地下根。高、中水位池杉细根的Fe和Mn浓度显著高于低水位, 其中Fe的浓度相差10倍以上, 但是叶的Fe、Mn浓度在不同水位之间没有显著差异。膝根的呼吸具有明显的季节差异, 8月和9月平均每个膝根的呼吸速率为2.1～2.5 mgCO₂&#8226;h^–1, 6月和11月为0.7～0.9 mgCO₂&#8226;h^–1, 3月为0.4 mgCO₂&#8226;h^–1; 膝根吸收O₂的摩尔数是释放CO₂摩尔数的4.6倍, 说明膝根吸收的O₂除了供自身呼吸, 大部分是提供给地下根利用。池杉之所以具有较强的耐水性, 与其在缺氧环境中能形成气生根和膝根、树干基部膨大和根系容重降低等有利于改善根系通气条件的生态适应机制密切相关。     

树木根系碳分配格局及其影响因子

根系作为树木提供养分和水分的“源”和消耗C的“汇”,在陆地生态系统C平衡研究中具有重要的理论意义。尽管20多年来的研究已经认识到根系消耗净初级生产力占总净初级生产力较大的比例,但是,根系（尤其是细根）消耗C的机理以及C分配的去向一直没有研究清楚。主要原因是细根消耗光合产物的生理生态过程相当复杂,准确估计各个组分消耗的C具有很大的不确定性,常常受树种和环境空间和时间异质性、以及研究方法的限制。综述了分配到地下的C主要去向,即细根生产和周转、呼吸及养分吸收与同化、分泌有机物、土壤植食动物,及有关林木地下碳分配机理的几种假说,分析了地下碳分配估计中存在的不确定性。目的是在全球变化C循环研究中对生态系统地下部分根系消耗的C以及分配格局引起重视。     

A plant root system architectural taxonomy: A framework for root nomenclature

Abstract

Research into root system morphology over the last two centuries has developed a diverse set of terminologies that are difficult to apply consistently across species and research specialties. In response to a need for better communication, a workshop held by the International Society for Root Research established some nomenclature standards for root research. These standards and their justification are presented in this study. A framework for a root system architectural taxonomy is created by defining four main classes of root: the tap root, that is, the first root to emerge from the seed; lateral roots, which are branches of other roots; shoot‐borne roots, which arise from shoot tissues; and basal roots, which develop from the hypocotyl, that is, the organ which is between the base of the shoot and the base of the tap root. It is concluded that adherence to the presented taxonomy will reduce confusion and eliminate some of the current confounding of results.     

模拟氮沉降对杉木幼苗细根的生理生态影响

细根对氮沉降的生理生态响应将显著影响森林生态系统的生产力和碳吸存。为了揭示氮沉降对杉木细根的生理生态影响,对一年生杉木(Cunninghamia lanceolata)幼苗进行了模拟氮沉降试验,并测定施氮1年后杉木幼苗细根生物量、细根形态学特征(比根长、比表面积)、元素化学计量学指标(C、N、P、C/N、C/P、N/P)、细根代谢特征(细根比呼吸速率、非结构性碳水化合物)。结果表明:(1)杉木细根生物量随氮添加水平的升高而显著降低,尤其是0—1 mm细根生物量;细根比根长和比表面积随氮添加水平升高而显著增大。(2)氮添加后杉木细根C含量、C/N、C/P显著降低,高氮添加导致1—2 mm细根N含量和N/P显著升高,而低氮添加导致1—2 mm细根P含量显著升高、N/P显著降低,而0—1 mm细根的N、P含量则保持相对稳定。(3)氮添加后杉木细根比呼吸速率无显著变化,细根可溶性糖含量随氮添加增加而显著增加,而淀粉含量和NSC显著降低。综合以上结果表明:氮添加后用于细根形态构建的碳分配减少,这可能会减少土壤中有机碳的保留,0—1 mm细根的形态更易发生变化,但是其内部N、P养分含量相对更稳定以维持生理活动,细根NSC对氮添加的响应表明施氮可能导致细根受光合产物的限制。     

Root turnover of groundnut (Arachis hypogaea L.) in soil tubes

Five groundnut cultivars were grown in transparent tubes of pasteurized loam compost in growth-chamber conditions. Weekly tracings were made of all the roots visible through the walls of the tubes. White roots were assessed as living, and brown or decayed roots as dead; this correlated with microscopical assessments of root viability based on cytoplasmic staining with neutral red followed by plasmolysis.For all five cultivars, root laterals began to die 3–4 weeks after plants were sown. Death of root laterals progressed down the soil profile with time, while new roots were produced successively deeper from the extending taproot. The half-life of individual roots was calculated as 3.7–4.4 weeks for all cultivars, based on assessments of the roots that died up to plant maturity (14–20 weeks, depending on cultivar). At maturity, 73–83% of the cumulative length of root systems had died. The onset and rate of root death were not related to onset of flowering or pod-filling; instead, the peak times of root death at different distances down the root system were related to earlier (3–5 week) peak times of root production in those regions. The net result of root turnover was that, despite continued new root production, the maximum length of living (white) roots of each cultivar was recorded at 2–4 weeks after sowing. Death of the earliest formed root laterals was also observed in the first five weeks after sowing of groundnut in an experimental field plot in Malawi. Progressive root turnover is considered to be a normal feature of groundnut, perhaps representing an energy-economy strategy.     

连作杨树人工林细根寿命的代际差异及其影响因素

细根寿命是调控森林生产力形成的关键。通过在连作Ⅰ、Ⅱ代杨树人工林固定样地内埋设微根管,对杨树不同根序细根年度生长动态开展连续观测并进行生存分析。结果表明,杨树不同根序细根累积生存率存在显著差异,高级根(3—5级)寿命较长,其累积生存率显著高于1级和2级细根。杨树细根寿命存在显著的代际差异,连作Ⅱ代人工林活根量、死根量和细根总量均高于Ⅰ代林。连作Ⅱ代人工林细根中位值寿命为(90±16)d,显著低于Ⅰ代人工林((102±22)d)。连作Ⅱ代林各根序细根数量、分布比例均高于Ⅰ代林,低级细根累积生存率低于Ⅰ代林而高级细根累积生存率显著高于Ⅰ代林。连作杨树人工林细根寿命显著受制于土壤环境,1级细根寿命与土壤速效氮相关性极显著(r=-0.861),2级细根寿命与土壤物理性状相关性较强且与土壤酚酸含量呈现极显著相关(r=0.870),高级根序细根寿命与土壤物理性质和养分状况等也具有一定相关性。连作杨树人工林土壤酚酸累积和养分有效性下降影响了细根寿命和周转,并进而造成净初级生产力损耗,相关结论为连作杨树人工林生产力衰退机理模型的建立提供了科学依据。     

Importance of internal hydraulic redistribution for prolonging the lifespan of roots in dry soil

Redistribution of water within plants could mitigate drought stress of roots in zones of low soil moisture. Plant internal redistribution of water from regions of high soil moisture to roots in dry soil occurs during periods of low evaporative demand. Using minirhizotrons, we observed similar lifespans of roots in wet and dry soil for the grapevine 'Merlot' (Vitis vinifera) on the rootstock 101-14 Millardet de Gramanet (Vitis riparia x Vitis rupestris) in a Napa County, California vineyard. We hypothesized that hydraulic redistribution would prevent an appreciable reduction in root water potential and would contribute to prolonged root survivorship in dry soil zones. In a greenhouse study that tested this hypothesis, grapevine root systems were divided using split pots and were grown for 6 months. With thermocouple psychrometers, we measured water potentials of roots of the same plant in both wet and dry soil under three treatments: control (C), 24 h light + supplemental water (LW) and 24 h light only (L). Similar to the field results, roots in the dry side of split pots had similar survivorship as roots in the wet side of the split pots (P = 0.136) in the C treatment. In contrast, reduced root survivorship was directly associated with plants in which hydraulic redistribution was experimentally reduced by 24 h light. Dry-side roots of plants in the LW treatment lived half as long as the roots in the wet soil despite being provided with supplemental water (P < 0.0004). Additionally, pre-dawn water potentials of roots in dry soil under 24 h of illumination (L and LW) exhibited values nearly twice as negative as those of C plants (P = 0.034). Estimates of root membrane integrity using electrolyte leakage were consistent with patterns of root survivorship. Plants in which nocturnal hydraulic redistribution was reduced exhibited more than twice the amount of electrolyte leakage in dry roots compared to those in wet soil of the same plant. Our study demonstrates that besides a number of ecological advantages to protecting tissues against desiccation, internal hydraulic redistribution of water is a mechanism consistent with extended root survivorship in dry soils.     

Effect of a Treatment with Naphthaleneacetic Acid on the Arrangement of Lateral Roots in Onion (Allium cepa L): General Pattern and Coordination between Ranks

Lateral roots in Allium cepa arise in longitudinal rows opposite the protoxylem poles of adventitious roots. The number, spacing within ranks and positional relationship to laterals in different ranks were analysed in control roots and those treated with an auxin (α-naphthaleneacetic acid). Treatment increased numbers of laterals per rank and distributed the more evenly in all ranks. Laterals of control roots were concentrated in two neighbouring ranks. Auxin-treated roots showed a more regular distribution of laterals between ranks and close spacing of laterals along each rank. Comparisons with theoretical random distributions suggest a dispersed spacing model for lateral root arrangement with mutual repulsion between successive lateral roots within each rank both in control and auxintreated roots. On the other hand, there is some interaction between laterals in different ranks in 0.1 mM NAA-treated roots.     

Root growth of wetland plants with different root types

A wastewater culture system was designed to study the root growth of eight species of wetland plants with two different root types. The system included a plastic barrel for holding the wastewater and a foam plate for holding the plant. The results indicated that the root growth of the plants with fibril roots was faster than that of the plants with rhizomatic roots. The species with fibril roots had higher root number (1349 per plant) than species with rhizomatic roots (549 per plant) after ten weeks of cultivation. The average root biomass of plants with fibril roots was 11.3 g per plant, whereas that of plants with rhizomatic roots was 7.4 g per plant. Fine root biomass of diameter ≤ 1 mm constituted 51.9% of the total root biomass in plants with fibril roots, whereas it accounted for only 25.1% in plants with rhizomatic roots. The root surface area of the plants with fibril roots (6933 cm² per plant) was markedly larger than that of the species with rhizomatic roots (1897 cm² per plant). The species with rhizomatic roots showed a longer root lifespan (46.6 days) than those with fibril roots (34.8 days).     

Temperature-induced alteration to root longevity in Lolium perenne

The production and mortality of roots is an important factor determining nutrient and carbon fluxes in terrestrial ecosystems. However, the effect of temperature on the longevity of roots is not understood. In this study the impact of changes in temperature on the longevity of Lolium perenne roots was determined.Plants were grown at 15 °C, 21 °C and 27 °C and their roots observed using a minirhizotron system. Major effects of temperature were measured with large reductions in root longevity occurring at higher temperatures. The implications of these data are discussed.     

落叶松和水曲柳不同根序细根形态结构、组织氮浓度与根呼吸的关系

根系具有高度的形态和生理功能异质性,在森林生态系统碳和养分循环中起重要作用。根系分枝的顺序构成根序,是根系最基本的构型特征,根序代表根系不同的发育阶段。然而,目前直接测定不同根序细根生理功能的研究很少。以落叶松（Larix gmelinii）和水曲柳（Fraxinus mandshurica）的细根为研究对象,使用气相氧电极测定不同根序细根的呼吸速率,探讨根系呼吸速率与其形态、结构和组织氮浓度的关系。结果表明：落叶松和水曲柳细根的直径、根长和维管束直径均随着根序的增加（1–5级）而增加,而比根长、组织氮浓度和呼吸速率随着根序的增加而降低,各根序之间差异显著（P〈0.05）;1级根比根长最大、皮层组织发达、组织氮浓度最高且呼吸速率也最高,其呼吸速率分别为17.57nmolO2·g^–1·s^–1（落叶松）和18.80 nmolO2·g^–1·s^–1（水曲柳）,比5级根分别高148%（落叶松）和124%（水曲柳）;并且,落叶松根的呼吸速率几乎有96%与根系组织氮浓度相关,而水曲柳根的呼吸速率则有89%与根系组织氮浓度相关。上述结果说明,细根的形态和生理功能异质性是紧密相连的,低级根的形态、结构决定其功能是吸收养分和水,而高级根的形态、结构决定其功能是运输和贮存养分。