贵州苏铁根的解剖学研究

对贵州苏铁（Cycas guizhouensis K.M.Lan et R.F.Zou）的根进行了解剖学研究,结果表明：（1）据外部形态和内部结构不同,贵州苏铁的根可分为正常根、珊瑚状根和肉质根三种类型。（2）正常根与肉质根的木栓形成层起源于表皮内方第1～2层皮层薄壁细胞,珊瑚状根的木栓形成层则起源于外方皮层的第2～3层细胞,而非中柱鞘。（3）首次发现肉质根一种新的增粗方式,是由周皮内方的皮层薄壁细胞恢复分裂能力,持续或周期性产生新的皮层薄壁细胞,引起根的增粗。     

贵州苏铁根的解剖学研究*

对贵州苏铁( Cycas guizhouensis K. M. Lan et R. F. Zou) 的根进行了解剖学研究, 结果表明: ( 1)
据外部形态和内部结构不同, 贵州苏铁的根可分为正常根、珊瑚状根和肉质根三种类型。( 2) 正常根与肉
质根的木栓形成层起源于表皮内方第1~ 2 层皮层薄壁细胞, 珊瑚状根的木栓形成层则起源于外方皮层的
第2~ 3 层细胞, 而非中柱鞘。( 3) 首次发现肉质根一种新的增粗方式, 是由周皮内方的皮层薄壁细胞恢
复分裂能力, 持续或周期性产生新的皮层薄壁细胞, 引起根的增粗。     

Root cap removal increases root penetration resistance in maize (Zea mays L)

The root cap assists the passage of the root through soil by means of its slimy mucilage secretion and by the sloughing of its outer cells. The root penetration resistance of decapped primary roots of maize (Zea mays L. cv. Mephisto) was compared with that of intact roots in loose (dry bulk density 1.0 g cm-3; penetration resistance 0.06 MPa) and compact soil (1.4 g cm-3; penetration resistance 1.0 MPa), to evaluate the contribution of the cap to decreasing the impedance to root growth. Root elongation rate and diameter were the same for decapped and intact roots when the plants were grown in loose soil. In compacted soil, however, the elongation rate of decapped roots was only about half that of intact roots, whilst the diameter was 30% larger. Root penetration resistances of intact and decapped seminal axis were 0.31 and 0.52 MPa, respectively, when the roots were grown in compacted soil. These results indicated that the presence of a root cap alleviates much of the mechanical impedance to root penetration, and enables roots to grow faster in compacted soils.     

Root cap influences root colonisation by Pseudomonas fluorescens SBW25 on maize

We investigated the influence of root border cells on the colonisation of seedling Zea mays roots by Pseudomonas fluorescens SBW25 in sandy loam soil packed at two dry bulk densities. Numbers of colony forming units (CFU) were counted on sequential sections of root for intact and decapped inoculated roots grown in loose (1.0 mg m(-3)) and compacted (1.3 mg m(-3)) soil. After two days of root growth, the numbers of P. fluorescens (CFU cm(-1)) were highest on the section of root just below the seed with progressively fewer bacteria near the tip, irrespective of density. The decapped roots had significantly more colonies of P. fluorescens at the tip compared with the intact roots: approximately 100-fold more in the loose and 30-fold more in the compact soil. In addition, confocal images of the root tips grown in agar showed that P. fluorescens could only be detected on the tips of the decapped roots. These results indicated that border cells, and their associated mucilage, prevented complete colonization of the root tip by the biocontrol agent P. fluorescens, possibly by acting as a disposable surface or sheath around the cap.     

树木不同着生位置1级根的形态、解剖结构和氮含量

树木根系中1级根在养分和水分吸收方面发挥着重要作用。研究1级根的形态结构与功能的联系,对了解1级根的生理功能和寿命,以及森林生态系统碳和养分的循环具有重要的理论意义。但是,1级根在根系统中,因着生的位置不同,可能表现出不同的生理生态功能。该研究以胡桃楸(Juglans mandshurica)、黄波罗(Phellodendron amurense)和水曲柳(Fraxinus mandshurica)人工林根系的1级根为研究对象,研究了不同着生位置的1级根的形态、解剖结构和组织化学特征。将1级根按着生位置的不同分成3类:Ar类根为2级根上的1级根;Br类根为2级–4级根的根尖;Cr类根为3级–5级根上的1级根。结果表明:不同着生位置的1级根,形态、解剖结构和组织化学方面都具有高度的异质性。3类1级根中,Ar类根数量多、根长较短、直径细,N含量高,皮层比例高、维根比低,主要由二原型原生木质部的根组成;Br类和Cr类根数量较少、单根较长、直径粗,N含量低,皮层比例低、维根比高,主要由多原型原生木质部的根组成。研究结果对了解不同着生位置的1级根的吸收功能和寿命具有重要的理论意义。     

Contribution of root cap mucilage and presence of an intact root cap in maize (Zea mays) to the reduction of soil mechanical impedance

BACKGROUND AND AIMS: The impedance to root growth imposed by soil can be decreased by both mucilage secretion and the sloughing of border cells from the root cap. The aim of this study is to quantify the contribution of these two factors for maize root growth in compact soil. METHODS: These effects were evaluated by assessing growth after removing both mucilage (treatment I -- intact) and the root cap (treatment D -- decapped) from the root tip, and then by adding back 2 micro L of mucilage to both intact (treatment IM -- intact plus mucilage) and decapped (treatment DM -- decapped plus mucilage) roots. Roots were grown in either loose (0.9 Mg m(-3)) or compact (1.5 Mg m(-3)) loamy sand soils. Also examined were the effects of decapping on root penetration resistance at three soil bulk densities (1.3, 1.4 and 1.5 Mg m(-3)). KEY RESULTS: In treatment I, mucilage was visible 12 h after transplanting to the compact soil. The decapping and mucilage treatments affected neither the root elongation nor the root widening rates when the plants were grown in loose soil for 12 h. Root growth pressures of seminal axes in D, DM, I and IM treatments were 0.328, 0.288, 0.272 and 0.222 MPa, respectively, when the roots were grown in compact soil (1.5 Mg m(-3) density; 1.59 MPa penetrometer resistance). CONCLUSIONS: The contributions of mucilage and presence of the intact root cap without mucilage to the lubricating effect of root cap (percentage decrease in root penetration resistance caused by decapping) were 43 % and 58 %, respectively. The lubricating effect of the root cap was about 30 % and unaffected by the degree of soil compaction (for penetrometer resistances of 0.52, 1.20 and 1.59 MPa).     

A Soil-free Plant Growth System to Facilitate Analysis of Plant Pathogen Interactions in Roots

Analysis of the interaction of pathogens with plant roots is often complicated by the growth of plants in a soil substrate. A soil-free plant growth system (SPS) was developed that removes the need for a substrate while supporting the growth of seedlings in a nutrient rich, oxygenated environment. The model legume Lupinus angustifolius was used to compare the growth of seedlings within soil and the SPS. Seedlings grown under both conditions were similar in morphology, anatomy and health (measured by leaf chlorophyll abundance) and importantly there was little difference in root growth and development although straighter and fuller root systems were achieved in the SPS. The ease of access to the root system proved efficient for the analysis of root and pathogen interactions with no interference from soil or adhering particulate matter. Following inoculation of L. angustifolius roots with Phytophthora cinnamomi the host/pathogen interaction was easily observed and tissues sampled undamaged.     

The position of localized soil compaction determines root and subsequent shoot growth responses

Plants growing in soils typically experience a mixture of loose and compact soil. The hypothesis that the proportion of a root system exposed to compact soil and/or the timing at which this exposure occurs determines shoot growth responses was tested. Broccoli (Brassica oleracea var. italica cv. Greenbelt) seedlings were grown in pot experiments with compact, loose and localized soil compaction created by either horizontal (compact subsoils 75 or 150 mm below loose topsoil) or vertical (adjacent compact and loose columns of soil) configurations of loose (1.2 Mg m(-3)) and compact (1.8 Mg m(-3)) soil. Entirely compact soil reduced leaf area by up to 54%, relative to loose soil. When compaction was localized, only the vertical columns of compact and loose soil reduced leaf area (by 30%). Neither the proportion of roots in compact soil nor the timing of exposure could explain the differing shoot growth responses to localized soil compaction. Instead, the strong relationship between total root length and leaf area (r(2)=0.92) indicated that localized soil compaction reduced shoot growth only when it suppressed total root length. This occurred when isolated root axes of the same plant were exposed to vertical columns of compact and loose soil. When a single root axis grew through loose soil into either a shallow or deep compact subsoil, compensatory root growth in the loose soil maintained total root length and thus shoot growth was unaffected. These contrasting root systems responses to localized soil compaction may explain the variable shoot growth responses observed under heterogeneous conditions.     

Field evaluation of laboratory techniques for predicting the ability of roots to penetrate strong soil and of the influence of roots on water sorptivity

The ability of two laboratory screening techniques to predict the abilities of roots of eight crop species to penetrate a compacted soil were evaluated and compared in a field experiment. A soil tilled to remove the effects of mechanical resistance was planted with the same species to serve as a control. Depth of root penetration, root density and the influence of the roots on the sorptivity of water were measured.Roots of all species penetrated deeper in the deep tilled than compacted soils. There were differences in the ability of roots of the species to penetrate the compacted soil. Generally dicotyledonous species had more roots penetrating to depth in both the compact and deep tilled soils. Within the main species classifications, lupin and safflower (dicotyledons) and oats and barley (monocotyledons) had the highest penetration into the compacted soil.Water sorptivities in the deep tilled soils were higher than those of the compact soil. Soil from planted treatments had higher sorptivities than soil which had not been planted. This is attributed to biopores left by the roots. Sorptivities of soils which had dicotyledonous species were generally higher than those of monocotyledons. The soil planted with safflower produced the highest sorptivity in the compacted layer (0.1–0.3 m).A comparison of the accuracy of the two laboratory screening methods in predicting the field penetration of roots suggest that the method involving mechanical stress was better than that involving osmotic stress. Relative root diameter was found to be a better indicator of the penetration ability of roots than relative root elongation.     

Soil strength influences wheat root interactions with soil macropores

Deep rooting is critical for access to water and nutrients found in subsoil. However, damage to soil structure and the natural increase in soil strength with depth, often impedes root penetration. Evidence suggests that roots use macropores (soil cavities greater than 75 μm) to bypass strong soil layers. If roots have to exploit structures, a key trait conferring deep rooting will be the ability to locate existing pore networks; a trait called trematotropism. In this study, artificial macropores were created in repacked soil columns at bulk densities of 1.6 g cm⁻³ and 1.2 g cm⁻³, representing compact and loose soil. Near isogenic lines of wheat, Rht-B1a and Rht-B1c, were planted and root–macropore interactions were visualized and quantified using X-ray computed tomography. In compact soil, 68.8% of root–macropore interactions resulted in pore colonization, compared with 12.5% in loose soil. Changes in root growth trajectory following pore interaction were also quantified, with 21.0% of roots changing direction (±3°) in loose soil compared with 76.0% in compact soil. These results indicate that colonization of macropores is an important strategy of wheat roots in compacted subsoil. Management practices to reduce subsoil compaction and encourage macropore formation could offer significant advantage in helping wheat roots penetrate deeper into subsoil.     

Root attributes affecting water uptake of rice (Oryza sativa) under drought

Lowland rice roots have a unique physiological response to drought because of their adaptation to flooded soil. Rice root attributes that facilitate growth under flooded conditions may affect rice response to drought, but the relative roles of root structural and functional characteristics for water uptake under drought in rice are not known. Morphological, anatomical, biochemical, and molecular attributes of soil-grown rice roots were measured to investigate the genotypic variability and genotype×environment interactions of water uptake under variable soil water regimes. Drought-resistant genotypes had the lowest night-time bleeding rates of sap from the root system in the field. Diurnal fluctuation predominated as the strongest source of variation for bleeding rates in the field and root hydraulic conductivity (Lp (r)) in the greenhouse, and was related to expression trends of various PIP and TIP aquaporins. Root anatomy was generally more responsive to drought treatments in drought-resistant genotypes. Suberization and compaction of sclerenchyma layer cells decreased under drought, whereas suberization of the endodermis increased, suggesting differential roles of these two cell layers for the retention of oxygen under flooded conditions (sclerenchyma layer) and retention of water under drought (endodermis). The results of this study point to the genetic variability in responsiveness to drought of rice roots in terms of morphology, anatomy, and function.     

Anatomical characteristics of individual roots within the fine-root architecture of Chamaecyparis obtusa (Sieb. & Zucc.) in organic and mineral soil layers

Nutrient availability and temporal variation of physical stress are usually higher in organic soil layers than in mineral soils. Individual roots within the fine-root system adjust anatomical, morphological, and turnover characteristics to soil conditions, for example nutrient availability and physical stresses. We investigated anatomical traits, including cork formation and passage and protoxylem cell numbers, in cross-sections of individual fine roots of the conifer Chamaecyparis obtusa (Siebold & Zucc.) growing under different soil conditions. The fine-root systems in different soil layers were compared by sampling ingrowth cores buried for 1 year and filled with organic and mineral soil substrates. The number of exodermal passage cells was lower in roots from organic soils than in those from mineral soils, suggesting that apical roots tend to be more stress-tolerant in the organic layer than in mineral soils. In contrast, both root tip and specific root tip density were higher in roots from organic soils than in those from mineral soil layers. The proportion of roots with two strands of protoxylem (diarch) was greater in organic (90%) than in mineral (25%) soils. Thus, although the absorptivity of individual apical roots decreases in organic layers, the absorptivity of the entire fine-root system of C. obtusa may be increased as a result of the increase in apical root density and the proportion of ephemeral roots. We found that the fine-root system had simultaneous plasticity in density, anatomy, and architecture in response to complex soil conditions.     

Scanning electron microscopic aspects of short tuberized roots, with special reference to cell rhizodermis evolution under drought and rehydration

Abstract. Different morphological aspects of short tuberized roots initiated during drought stress in mesophytic species, such as Sinapis alba L., were studied with the scanning electron microscopy technique of cryofixation. Specially adapted for direct and immediate observation of fresh living roots, this method has given precise information about rhizodermal organization and evolution during drought and rehydration.
The main difference from a normal lateral root grown in a well-watered soil appears in the basal enlarged zone of the short root where all the cells show the same round and turgid aspect. In the medium zone of the short root, rhizodermis differentiation into alternating rows of short (trichoblasts) and long (atrichoblasts) cells, which characterizes the typical Sinapis root, remains clearly discernible, though not so regular as in normal lateral roots.
The turgid state of rhizodermal cells all over the short tuberized root grown in a drying soil suggests an effective regulatory mechanism for water deficit avoidance.
During the first hours of rehydration, immediate absorption of water is noticeable through the rapid swelling of some long cells which appear to protrude considerably among other rhizodermal cells. However, these protrusions will not give rise to hairs, as further observations of short roots after growth has resumed show very distinctly that progressive hair formation occurs in the medium zone of the root, and that their emergence originates from trichoblasts only.
These observations may indicate that atrichoblasts, on account of their highly vacuolated condition, are the first cells to absorb water and that they may even be stimulated, in some environmental conditions, to initiate hair formation, although they are not so well adapted to do so as the short cells in this species.     

Soil physical strength rather than excess ethylene reduces root elongation of Eucalyptus seedlings in mechanically impeded sandy soils

Seedling establishment in heavily compact soils is hampered by poor root growth caused by soil chemical or physical factors. This study aims to determine the role of ethylene in regulating root elongation through mechanically impeded sandy soils using Eucalyptus todtiana F. Muell seedlings. Concentrations of ethephon (1, 10, and 100???M) were added to non-compact soils, and endogenous ethylene production from seedling roots was compared to ethylene production of roots grown in physically compacted field soils (98.6?% sand). The ethylene-inhibitor 3,5-diiodo-4-hydroxybenzoic acid (DIHB) (0.1???M) was included for each treatment to counteract the negative effects of excess ethylene or compact soils on root elongation. Root elongation was reduced in high ethylene soils by 49?% and high bulk density soils by 44?%. Root ethylene production increased ninefold in roots grown in the high ethylene environment (100???M), but decreased 80?% in compact soils. The use of DIHB did not alter root length and produced varying results with respect to ethylene production, suggesting an interaction effect involving high amounts of soil ethylene. While ethylene regulates root growth, the physical strength of sandy soils is the major factor limiting root elongation in mechanically impeded soils.     

The Developmental Anatomy of the Root System in Yam Bean, Pachyrhizus erosus Urban

Investigations revealed that the anatomy of the primary radicularroot of yam bean (Pachyrhizus erosus L.) was typically dicotyledonousexcept that the xylem was not completely developed centripetally.Most of the roots had tetrarch xylem, although a few triarchand pentarch roots were also observed. In both tuberous andnon-tuberous roots, secondary thickening occurred by the formationof the meristematic vascular cambium which formed secondarytissues in a normal fashion. Subsequently, tuberization wasinitiated in the secondary xylem by the development of anomalous‘secondary’ cambia from parenchyma cells surroundingvessel elements. Anomalous ‘secondary’ cambia alsodeveloped from parenchyma cells not associated with vessels.Subsequently, anomalous ‘tertiary’ cambia differentiatedfrom tissues produced by the anomalous ‘secondary’cambia. Activities of these anomalous cambia resulted in theproduction of parenchyma storage cells and were chiefly responsiblefor the growth of the mature tuber. Pachyrhizus erosus L., yam bean, tuberous root, anatomy, anomalous ‘secondary’ cambia, anomalous ‘tertiary’ cambia, centripetal xylem development     

Introducing short roots in a desert perennial: anatomy and spatiotemporal foraging responses to increased precipitation

? The desert flora possesses diverse root architectures that result in fast growth in response to precipitation. We introduce the short root, a previously undescribed second-order root in the aridland chamaephyte Cryptantha flava, and explore fine root production. ? We describe the short root anatomy and associated fine roots, correlate standing fine root crop with soil moisture, and explore the architectural level - the short root, third-order lateral roots, or the whole root system - at which fine roots are induced by watering and the amount of water required. ? We show that short roots are borne at intervals on lateral roots and produce fine roots at their tips; new fine roots are white and have root hairs, while brown and black fine roots are apparently dead; and fine root production is triggered at the level of lateral roots and with relatively low precipitation (≤ 2 cm). ? Short roots are suberized and thus are probably not capable of water uptake themselves, but serve as initiation sites for fine roots that grow rapidly in response to rainfall. Thus, C. flava should be a beneficiary of projected precipitation increases in habitats where rainfall is pulsed.     

Root plasticity under abiotic stress

Abiotic stresses increasingly threaten existing ecological and agricultural systems across the globe. Plant roots perceive these stresses in the soil and adapt their architecture accordingly. This review provides insights into recent discoveries showing the importance of root system architecture (RSA) and plasticity for the survival and development of plants under heat, cold, drought, salt, and flooding stress. In addition, we review the molecular regulation and hormonal pathways involved in controlling RSA plasticity, main root growth, branching and lateral root growth, root hair development, and formation of adventitious roots. Several stresses affect root anatomy by causing aerenchyma formation, lignin and suberin deposition, and Casparian strip modulation. Roots can also actively grow toward favorable soil conditions and avoid environments detrimental to their development. Recent advances in understanding the cellular mechanisms behind these different root tropisms are discussed. Understanding root plasticity will be instrumental for the development of crops that are resilient in the face of abiotic stress.

Recent discoveries show the importance of root system architecture plasticity for the survival and growth of plants under several abiotic stresses.     

The impact of different soil moisture and soil compaction on the growth of triticale root system

Effects of different soil moisture (soil drought and waterlogging) and soil compaction (1.33 and 1.50 g·cm⁻³) on the growth and morphological traits of the root system were studied in four breeding forms and seven cultivars of triticale. Morphological changes, including the restriction of root extension, expansion and proliferation of laterals roots, occur in plants grown in different soil moisture and in compact soil. The investigations comprised quantitative and qualitative analyses of a developed plant root system through determining the number, length and dry matter of the particular components of the root system. Obtained results have demonstrated a relatively broad variation in the habit of the triticale root system. Plants grown under compact soil and low or high soil water content showed a smaller number and less dry matter of lateral branching than plants grown in control conditions. The harmful effects of compact soil and drought conditions on the growth of roots was greater when compared with that of plants exposed to waterlogging. The observed effects of all treatments were more distinct in a drought sensitive strains. The drought resistant forms were a more characterize with extensive rooting and by smaller alterations in the root morphology under the stress conditions compared with drought sensitive one. Results confirm that the breeding forms (CHD-12 and CHD-173) of a high drought susceptibility was found to be also more sensitive to periodical soil water excess. A more efficient water use and a lower shoot to root (S/R) ratio were found to be major reasons for a higher stress resistance of the breeding forms (CHD-220 and CHD-247). The reasons for a different response of the examined breeding forms and cultivars to the conditions of drought or waterlogging may be a more economical water balance and more favourable relations between the shoot and root dimensions in the drought resistant forms and cultivars. The results suggest that the morphological traits of the triticale root system may be used in practice as direct or indirect selection criteria in maize breeding.     

Anatomy and Histochemistry of the Root System of the Kiwifruit Vine, Actinidia deliciosa var. deliciosa.

The kiwifruit vine is a species which has been newly introducedinto cultivation and little is known of its comparative physiologyand anatomy. In this study we found that fibrous, 'magnolioid'roots, which have undergone secondary vascular development butwhich retain the cortex and develop a suberized epidermis, comprisethe greater part of the root system (95% of total length). Newlyinitiated roots with primary development conform to norms establishedin other woody plant species. However, the structural roots,like the fibrous roots, also retain a cortex and phellodermwhich is initiated by hypodermal cells within the cortex andnot by the pericycle which is the common progenitor tissue inother species. This phellogen produces new cells centrifugallyonly. The cortex is a relatively small component of the structuralroot and the bulk of the tissue is vascular in origin, as inthe roots of other plant species. The endodermis is retainedand continues to divide periclinally to accommodate the increasein circumference with growth.Copyright 1993, 1999 Academic Press Actinidia deliciosa, root anatomy, ontogony, histochemistry, exodermis, endodermis