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.     

Numbers and locations of native bacteria on field-grown wheat roots quantified by fluorescence in situ hybridization (FISH)

Native bacteria, Pseudomonas and filamentous bacteria were quantified and localized on wheat roots grown in the field using fluorescence in situ hybridization (FISH). Seminal roots were sampled through the season from unploughed soil in a conservation farming system. Such soils are spatially heterogeneous, and many roots grow slowly through hard soil with cracks and pores containing dead roots remnant from previous crops. Root and rhizosphere morphology, and contact with soil particles were preserved, and autofluorescence was avoided by observing sections in the far-red with Cy5 and Cy5.5 fluorochromes. Spatial analyses showed that bacteria were embedded in a stable matrix (biofilm) within 11 microm of the root surface (range 2-30 microm) and were clustered on 40% of roots. Half the clusters co-located with axial grooves between epidermal cells, soil particles, cap cells or root hairs; the other half were not associated with visible features. Across all wheat roots, although variable, bacteria averaged 15.4 x 10(5) cells per mm(3) rhizosphere, and of these, Pseudomonas and filaments comprised 10% and 4%, respectively, with minor effects of sample time, and no effect of plant age. Root caps were most heavily colonized by bacteria along roots, and elongation zones least heavily colonized. Pseudomonas varied little with root development and were 17% of bacteria on the elongation zone. Filamentous bacteria were not found on the elongation zone. The most significant factor to rhizosphere populations along a wheat root, however, was contact with dead root remnants, where Pseudomonas were reduced but filaments increased to 57% of bacteria (P < 0.001). This corresponded with analyses of root remnants showing they were heavily colonized by bacteria, with 48% filaments (P < 0.001) and 1.4%Pseudomonas (P = 0.014). Efforts to manage rhizosphere bacteria for sustainable agricultural systems should continue to focus on root cap and mucilage chemistry, and remnant roots as sources of beneficial bacteria.     

亚热带典型树种根毛特征及其与共生真菌的关系

根毛和共生真菌增加了吸收面积,提高了植物获取磷等土壤资源的能力。由于野外原位观测根表微观结构较为困难,吸收细根、根毛、共生真菌如何相互作用并适应土壤资源供应,缺乏相应的数据和理论。该研究以受磷限制的亚热带森林为对象,选取了21种典型树种,定量了根毛存在情况、属性变异,分析了根毛形态特征与共生真菌侵染率、吸收细根功能属性之间的关系,探讨了根表结构对低磷土壤的响应和适应格局。结果表明:1)在亚热带森林根毛不是普遍存在的, 21个树种中仅发现7个树种存有根毛, 4个为丛枝菌根(AM)树种, 3个为外生菌根(ECM)树种。其中,马尾松(Pinus massoniana)根毛出现率最高,为86%;2)菌根类型是理解根-根毛-共生真菌关系的关键,AM树种根毛密度与共生真菌侵染率正相关,但ECM树种根毛直径与共生真菌侵染率负相关; 3) AM树种根毛长度和根毛直径、ECM树种根毛出现率与土壤有效磷含量呈负相关关系。该研究揭示了不同菌根类型树种根毛-共生真菌-根属性的格局及相互作用,为精细理解养分获取策略奠定了基础。     

Response of wheat to subsoil salinity and temporary water stress at different stages of the reproductive phase

To examine the effects of subsoil NaCl salinity in relation to water stress imposed at different growth stages, wheat was grown in a heavy texture clay soil (vertosol) under glasshouse conditions in polythene lined cylindrical PVC pots (100 cm long with 10.5 cm diameter) with very low salinity level (EC_e 1.0 dS/m; ESP 1.0 and Cl 30 mg/kg soil) in top 10 cm soil (10–20 cm pot zone) and low salinity level (EC_e 2.5 dS/m, ESP 5, and Cl 100 mg/kg soil) in top 10–20 cm soil (20–30 cm pot zone). The plants were exposed to three subsoil salinity levels in the 20–90 cm subsoil (30–100 cm pot zone) namely low salinity (EC_e: 2.5 dS/m, ESP: 5, Cl: 100 mg/kg soil), medium salinity (EC_e: 4.0 dS/m, ESP: 10, Cl: 400 mg/kg) and high salinity (EC_e: 11.5 dS/m, ESP: 20, Cl: 1950 mg/kg) in the subsoil (20–90 cm soil layer: 30–100 cm pot zone). Watering of plants was withheld for 20 days commencing at either early booting or anthesis or mid grain filling, and then resumed until maturity, and these treatments were compared with no water stress. Water stress commencing at anthesis stage had the most depressing effect on grain yield and water use efficiency of wheat followed by water stress at grain filling stage and early booting stage. High subsoil salinity reduced grain yield by 39.1, 24.3%, and 13.4% respectively in plants water-stressed around anthesis, early booting, and mid grain filling compared with 36.6% in well-watered plants. There was a significant reduction in root biomass, rooting depth, water uptake and water use efficiency of wheat with increasing subsoil salinity irrespective of water regimes. Plants at high subsoil salinity had 64% of their root biomass in the top 0–30 cm soil and there was a marked reduction in subsoil water uptake. Roots also penetrated below the non-saline surface into salinised subsoil and led to attain high concentration of Na and Cl and reduced Ca/Na and K/Na ratio of flag leaf at anthesis stage. Results suggest that high subsoil salinity affects root growth and water uptake, grain yield and water use efficiency even in well water plants. Water stress at anthesis stage had the most depressing effect on wheat.     

Drought-induced changes in soil contact and hydraulic conductivity for roots of Opuntia ficus-indica with and without rhizosheaths

Water movement between roots and soil can be limited by incomplete root–soil contact, such as that caused by air gaps due to root shrinkage, and can also be influenced by rhizosheaths, composed of soil particles bound together by root exudates and root hairs. The possible occurrence of air gaps between the roots and the soil and their consequences for the hydraulic conductivity of the root–soil pathway were therefore investigated for the cactus t Opuntia ficus-indica, which has two distinct root regions: a younger, distal region where rhizosheaths occur, and an older, proximal region where roots are bare. Resin-embedded sections of roots in soil were examined microscopically to determine root–soil contact for container-grown plants kept moist for 21 days, kept moist and vibrated to eliminate air gaps, droughted for 21 days, or droughted and vibrated. During drought, roots shrank radially by 30% and root–soil contact in the bare root region of nonvibrated containers was reduced from 81% to 31%. For the sheathed region, the hydraulic conductivity of the rhizosheath was the least limiting factor and the root hydraulic conductivity was the most limiting; for the bare root region, the hydraulic conductivity of the soil was the least limiting factor and the hydraulic conductivity of the root–soil air gap was the most limiting. The rhizosheath, by virtually eliminating root–soil air gaps, facilitated water uptake in moist soil. In the bare root region, the extremely low hydraulic conductivity of the root–soil air gap during drought helped limit water loss from roots to a drier soil.     

The effect of soil strength on the growth of pigeonpea radicles and seedlings

The effect of soil strength on the growth of pigeonpea radicles and seedlings was investigated in cores of three clay soils prepared at different water contents and bulk densities in the laboratory.Radicle elongation directly into soil cores was reduced from 50–70 mm d^-1 at strengths less than 0.5 MPa to 0 mm d^-1 at 3.5–3.7 MPa. The response to soil strength was affected by the water content of the soil, presumably as a result of reduced oxygen availability in wetter soil. This effect was apparent in soils wet to air-filled porosities less than 0.15 m³ m^-3.Radicles were more sensitive to high soil strength (>1.5 MPa) than were seedling roots which encountered the same conditions at 60 mm in the profile. Radicle growth ceased at 3.5 MPa which reduced seedling root growth by only 60%.Despite a 60% reduction in root length in the high strength zone, seedling roots compensated in zones of loose soil above and below the compacted layer, and total root length and shoot growth were unaffected. There was no evidence of a root signal response which results in reduced shoot growth in some species in response to high soil strength.The proliferation of roots in surface layers and the delayed penetration of the root system to depth in compacted soil are likely to expose seedlings to a greater risk of water-deficit in the field, particularly under dryland conditions where plants rely on stored subsoil water for growth.     

Root effects on soil water and hydraulic properties

Plants can affect soil moisture and the soil hydraulic properties both directly by root water uptake and indirectly by modifying the soil structure. Furthermore, water in plant roots is mostly neglected when studying soil hydraulic properties. In this contribution, we analyze effects of the moisture content inside roots as compared to bulk soil moisture contents and speculate on implications of non-capillary-bound root water for determination of soil moisture and calibration of soil hydraulic properties. In a field crop of maize (Zea mays) of 75 cm row spacing, we sampled the total soil volumes of 0.7 m × 0.4 m and 0.3 m deep plots at the time of tasseling. For each of the 84 soil cubes of 10 cm edge length, root mass and length as well as moisture content and soil bulk density were determined. Roots were separated in 3 size classes for which a mean root porosity of 0.82 was obtained from the relation between root dry mass density and root bulk density using pycnometers. The spatially distributed fractions of root water contents were compared with those of the water in capillary pores of the soil matrix. Water inside roots was mostly below 2–5% of total soil water content; however, locally near the plant rows it was up to 20%. The results suggest that soil moisture in roots should be separately considered. Upon drying, the relation between the soil and root water may change towards water remaining in roots. Relations depend especially on soil water retention properties, growth stages, and root distributions. Gravimetric soil water content measurement could be misleading and TDR probes providing an integrated signal are difficult to interpret. Root effects should be more intensively studied for improved field soil water balance calculations. Presented at the International Conference on Bioclimatology and Natural Hazards, Pol’ana nad Detvou, Slovakia, 17–20 September 2007.     

Soil structure and plant growth: Impact of bulk density and biopores

Compacted soils are not uniformly hard; they usually contain structural cracks and biopores, the continuous large pores that are formed by soil fauna and by roots of previous crops. Roots growing in compacted soils can traverse otherwise impenetrable soil by using biopores and cracks and thus gain access to a larger reservoir of water and nutrients. Experiments were conducted in a growth chamber to determine the plant response to a range of uniform soil densities, and the effect of artificial and naturally-formed biopores. Barley plants grew best at an intermediate bulk density, which presumably represented a compromise between soil which was soft enough to allow good root development but sufficiently compact to give good root-soil contact. Artificial 3.2 mm diameter biopores made in hard soil gave roots access to the full depth of the pot and were occupied by roots more frequently than expected by chance alone. This resulted in increased plant growth in experiments where the soil was allowed to dry. Our experiments suggest that large biopores were not a favourable environment for roots in wet soil; barley plants grew better in pots containing a network of narrow biopores made by lucerne and ryegrass roots, responded positively to biopores being filled with peat, and some pea radicles died in biopores. A theoretical analysis of water uptake gave little support to the hypothesis that water supply to the leaves was limiting in either very hard or very soft soil. The net effect of biopores to the plant would be the benefits of securing extra water and nutrients from depth, offset by problems related to poor root-soil contact in the biopore and impeded laterals in the compacted biopore walls.     

Direct evidence on participation of root hairs in phosphorus (32P) uptake from soil

Root hairs substantially extend root surface for ion uptake. Although many reports suggest a relationship between root hairs and phosphorus (P) uptake of plants, the role of root hairs in phosphorus uptake from soils is still debated. We measured uptake of phosphorus from soil directly via root hairs. Root hairs only were allowed to penetrate through a tightly stretched nylon screen (53 µm) glued to the bottom of a PVC tube. The penetrating root hairs grew for 2 and 4 days in soil labelled with radioisotope phosphorus (P) tracer ³²P (185 kBq g^-1 dry soil) filled in another PVC tube. Transparent plastic rings of thickness ranging from 0.25 mm to 2.0 mm were inserted between the two PVC tubes. This provided slit width for microscopic observations in situ, which confirmed that only root hairs were growing into the ³²P labelled soil. In some cases no rings were inserted (slit width = 0) where both root hairs and root surface were in contact with the labelled soil (total ³²P uptake). The uptake of³² P from soil via the root hairs only was quantified by measuring activity of ³²P in the plant shoot (³²P uptake only via root hairs).The results showed that when 70 percent of the root hairs grew into the labelled soil, they contributed to 63 percent of the total P uptake. With decreasing number of root hairs growing into the ³²P labelled soil, the quantity of ³²P in the plant shoot decreased. In this study, P uptake via root hairs was measured in a soil-based system, where root hairs were the only pathway of ³²P from soil to the plant shoot. Therefore, this study provides a strong evidence on the substantial participation of root hairs in uptake of phosphorus from soil.     

森林生态系统林木根系对优先流的影响

土壤水和溶质运移是土壤学和环境科学研究的难点和热点,优先流是一种常见的土壤溶质运移形式,绕过土壤基质而优先运移至地下水源,造成土壤养分的流失和水质的恶化。林木根系是土壤层的重要部分,其结构形态影响着优先流过程,为量化林木根系结构对土壤优先流的影响,以首都圈森林生态系统鹫峰定位监测站为研究区域,利用野外染色示踪与室内分析相结合的方法,定量分析根长密度和根系生物量在优先流区和基质流区的变化。结果表明:1)随着土层深度的增加,根长密度表现为减小的趋势,对径级d1 mm,1d3 mm和3d5 mm根系而言,根长密度在优先流区大于基质流区发生概率分别为66.7%,88.9%和83.3%;2)根系d1 mm对优先流贡献度最大,均值为94.8%,1d3 mm和3d5 mm根系对优先流贡献度较小,均值分别为4.3%和0.9%;3)研究点根系生物量进行统计,66.7%优先流区根系生物量大于基质流区根系生物量。开展根系对优先流的影响研究,有助于探明土壤水分运移规律,分析地表地下水质恶化根源,为生态环境安全提供理论指导和技术支持。     

The effects of free-air CO2 enrichment and soil water availability on spatial and seasonal patterns of wheat root growth

Spring wheat [ Triticum aestivum (L). cv. Yecora Rojo] was grown from December 1992 to May 1993 under two atmospheric CO₂ concentrations, 550 μmol mol^–1 for high-CO₂ plots, and 370 μmol mol^–1 for control plots, using a Free-Air CO₂ Enrichment (FACE) apparatus. In addition to the two levels of atmospheric CO₂, there were ample and limiting levels of water supply through a subsurface trip irrigation system in a strip, split-plot design. In order to examine the temporal and spatial root distribution, root cores were extracted at six growth stages during the season at in-row and inter-row positions using a soil core device (86 mm ID, 1.0 m length). Such information would help determine whether and to what extent root morphology is changed by alteration of two important factors, atmospheric CO₂ and soil water, in this agricultural ecosystem. Wheat root growth increased under elevated CO₂ conditions during all observed developmental stages. A maximum of 37% increase in total root dry mass in the FACE vs. Control plots was observed during the period of stem elongation. Greater root growth rates were calculated due to CO₂ enhancement until anthesis. During the early vegetative growth, root dry mass of the inter-row space was significantly higher for FACE than for Control treatments suggesting that elevated CO₂ promoted the production of first-order lateral roots per main axis. Then, during the reproductive period of growth, more branching of lateral roots in the FACE treatment occurred due to water stress. Significant higher root dry mass was measured in the inter-row space of the FACE plots where soil water supply was limiting. These sequential responses in root growth and morphology to elevated CO₂ and reduced soil water supports the hypothesis that plants grown in a high-CO₂ environment may better compensate soil-water-stress conditions.     

Biomass and distribution of fine and coarse roots from blue oak (Quercus douglasii) trees in the northern Sierra Nevada foothills of California

This research adds to the limited data on coarse and fine root biomass for blue oak (Quercus douglasii Hook and Arn.), a California deciduous oak species found extensively throughout the interior foothills surrounding the Central Valley. Root systems of six blue oak trees were analyzed using three methods — backhoe excavation, quantitative pits, and soil cores. Coarse root biomass ranged from 7 to 177 kg per tree. Rooting depth for the main root system ranged from 0.5 to 1.5 m, with an average of 70% of excavated root biomass located above 0.5 m. Of the total biomass in excavated central root systems, primary roots (including burls) accounted for 56% and large lateral roots (> 20 mm diameter) accounted for 36%. Data from cores indicated that most biomass outside of the root crown was located in fine roots and that fine root biomass decreased with depth. At surface depths (0–20 cm), small-fine (< 0.5 mm diameter) roots accounted for 71%, large-fine (0.5–2.0 mm) for 25%, and coarse (> 2 mm) for 4% of total root biomass collected with cores. Mean fine root biomass density in the top 50 cm was 0.43 kg m⁻³. Fine root biomass did not change with increasing distance from the trees (up to approximately 5 m). Thus, fine roots were not concentrated under the tree canopies. Our results emphasize the importance of the smallest size class of roots (<0.5 mm), which had both higher N concentration and, in the area outside the central root system, greater biomass than large fine (0.5–2.0 mm) or coarse (> 2.0 mm) roots. This revised version was published online in June 2006 with corrections to the Cover Date.     

不同水分条件对冬小麦根系时空分布、土壤水利用和产量的影响

为了明确华北严重缺水区晚播冬小麦灌水对根系时空分布和土壤水分利用规律的影响,以冬小麦石麦15为材料,利用田间定位试验研究了不同灌水处理(春季不灌水W0;春季灌拔节水75mm,W1;春季灌起身水、孕穗水和灌浆水共225mm,W3)对根系干重密度(DRWD)、根长密度(RLD)、体积密度、分枝数等在0—200cm土层的垂直分布、动态变化及其对耗水和产量的影响,结果表明:随着春季灌水量的减少,开花后0—80cm土层的根干重密度、根长度密度、体积密度和分枝数密度均显著减少,80cm—200m土层的根干重密度、根长度密度、体积密度和分枝数密度却显著增加,并且显著增加冬小麦在灌浆期间对100cm以下深层土层水分的利用,总耗水量W1和W0分别比W3减少70.9mm、115.1mm,土壤耗水量分别比W3增加79.1mm、108.9mm,子粒产量W1和W0分别比W3减少653.3kg/hm2、1470kg/hm2,水分利用效率(WUE)则分别比W3提高0.09kg/m3、0.06kg/m3。晚播冬小麦春季灌1水(拔节水)可以促进根系深扎,增加深土层的根系分布量,提高对深层土壤贮水的吸收利用量,有利于实现节水与高产的统一。     

Root clumping may affect the root water potential and the resistance to soil-root water transport

We have appraised for clumped root systems the widely-accepted view that the resistance to water flux from soil to roots (‘soil resistance’) is low under most field conditions, so that root water potential would closely follow the mean soil water potential. Three root spatial arrangements were studied, simulating either the regular pattern generally assumed in models, or two degrees of root clumping frequently observed in the field. We used a numerical 2-dimensional model of water transfer which assumes a control of evapotranspiration by root signalling. Calculations were carried out at two evaporative demands and for two contrasting soil hydraulic properties. The rate of soil depletion, the timing of the reduction in evapotranspiration and the difference between root water potential and mean soil water potential were all affected by the root spatial arrangement, with a greater effect at high evaporative demand and low soil hydraulic conductivity. Almost all the soil water reserve was available to plants without reduction in evapotranspiration in the regular case, while only a part of it was available in clumped cases. In the regular case, calculated ‘soil resistances’ were similar to those calculated using Newman's (1969) method. Conversely they were higher by up to two orders of magnitude in clumped root spatial arrangements. These results place doubt on the generality of the view that ‘soil resistance’ is low under common field conditions. They are consistent with the results of field experiments, especially with recent data dealing with root-to-shoot communication.     

Potential importance of the subsoil for the P and Mg nutrition of wheat

A method is described which allowed the quantification of the potential uptake of P and Mg from the subsoil (>30cm) by spring wheat. Wheat was grown on an artificial topsoil (sand with no plant available P or Mg) which was superimposed on loess subsoils in N. Germany. The supply of P and Mg in the topsoil was varied by application of different quantities of P and Mg fertilizer. Uptake of P and Mg from the subsoil was calculated as the difference between total plant uptake (determined by plant analysis) and the quantities of P and Mg removed from the topsoil (determined by soil analysis). P uptake from the subsoil increased from 37% to 85% of total P uptake, with decreasing P supply in the topsoil. Calculations of potential supply by diffusion showed that, with a CAL-extractable P₂O₅ content in the subsoil of 9 mg 100g^-1, supply from the subsoil was only possible if the influence of root hairs was considered. The method also showed that the total demand for Mg by spring wheat could be satisfield from the supply of Mg from the subsoil of typical loess soils. Mg uptake from the subsoil decreased to 33% of total uptake with increasing Mg supply in the topsoil.     

Root:soil adhesion in the maize rhizosphere: the rheological approach

This study was designed to investigate the strength of attachment of plant seedling roots to the soil in which they were grown. The study also assessed the effects of differing soil textures and differing soil matric potentials upon the strength of the root:soil attachment. A device for growing roots upon a soil surface was designed, and was used to produce roots which were attached to the soil. In order to quantify root:soil adhesion, roots of maize seedlings, grown on the soil surface, were subsequently peeled off using a universal test machine, in conjunction with simultaneous time-lapse video observation. To clarify the partitioning of energy in the root:soil peeling test, separate mechanical tests on roots, and on two adherent remoulded topsoil balls were also carried out. The seedling root was characterised by a low bending stiffness. The energy stored in bending was negligible, compared to the root:soil adhesion energy. The mechanical properties of two adherent remoulded topsoil balls were a decrease of the soil:soil adhesion energy as the soil:soil plastic energy increased. These two parameters were therefore interdependent. Using a video-camera system, it was possible to separate the different processes occurring during the root:soil peeling test, in particular, the seed:soil adhesion and the root:soil soil adhesion. An interpretation of the complex and variable force:displacement curves was thus possible, enabling calculation of the root:soil interfacial rupture energy. At a given suction (10 kPa), the results of the peeling test showed a clear soil texture effect on the value of the root:soil interfacial rupture energy. In contrast, for the same silty topsoil, the effect of the soil water suction on the value of the interfacial rupture energy was very moderate. The root:soil interfacial rupture energy was controlled mainly by a product of microscopic soil specific surface area and the macroscopic contact surface area between the root and the soil. Biological and physical interactions contributing to root:soil adhesion such as root:soil interlocking mechanics were also analysed and discussed. This revised version was published online in June 2006 with corrections to the Cover Date.     

Capacity of biological soil crusts colonized by the lichen <Emphasis Type="Italic">Diploschistes</Emphasis> to metabolize simple phenols

Background and aims

Soil compaction strongly affects water uptake by roots. The aim of the work was to examine soil—plant interactions with focus on the impact of distribution of compacted soil layers on growth and water uptake by wheat roots.

Methods

The growth-chamber experiment was conducted on wheat growth in soil with compacted soil layers. The system for maintaining constant soil water potential and measurement of daily water uptake from variously compacted soil layers was used.

Results

Layered soil compaction differentiated vertical root distribution to higher extent for root length than root mass. The propagation rate of a water extraction front was the highest through layers of moderately compacted soil. The root water uptake rate was on average 67 % higher from moderately than heavily compacted soil layers. Correlations between water uptake and the length of thick roots were increasing with increasing level of soil compaction.

Conclusions

The study shows that root amount, water uptake, propagation of water extraction and shoot growth strongly depend on the existence of compacted layers within soil profile. The negative effects of heavily compacted subsoil layer on water uptake were partly compensated by increased uptake from looser top soil layers and significant contribution of thicker roots in water uptake.     

Root growth of triticale and barley grown for grain or for forage-plus-grain in a Mediterranean climate

Root systems of one triticale (× triticosecale Witt.) and one barley (Hordeum vulgare L.) cultivar grown for grain or for the dual purpose of winter forage-plus-grain were studied in a Mediterranean climate (Granada, Spain). The aim was to assess the effect of winter forage removal on root systems and to improve the knowledge of cereal root systems under Mediterranean conditions in relation to soil water use. After the forage was removed by clipping at the end of the winter period, cereal roots were shallower and lower in length densities, compared to the unclipped treatment. The largest differences occurred during the clipping-anthesis period and in the upper soil layers. At the end of the life cycle, the differences between the two systems regarding depth, density and dry matter of roots were small or nil. Moreover, there were no differences in total water use between clipped and unclipped cereals.Under both production systems (grain and forage plus grain), cereals demonstrated variable downward root extension (0.9 to 1.8 m) as a response to the wetting depth. Triticale roots continued growing after anthesis, especially in the deeper soil layers. In spite of that, root systems were not able to extract a notable amount of residual water (25 to 50 mm) apparently available from the subsoil. In semi-arid Mediterranean drylands, cereal root systems with greater phenotypic plasticity (deeper or larger in the subsoil) in response to subsoil water should be of interest in wetter areas or seasons. This does not necessarily imply a larger root system, but rather a root growth pattern with greater root growth in the subsoil.     

土壤水分对黄土高原主要造林树种细根表面积季节动态的影响

在黄土丘陵沟壑区陕西省安塞县,于2007年生长季内,采用根钻法对刺槐(Robinia pseudoacacia)、侧柏(Platycladus orientalis)、油松(Pinus tabulaeformis)林地的细根和土壤水分进行了动态调查。结果表明:生长季内,刺槐、侧柏、油松林地0-200cm土层的土壤含水量变动较大,此土层是树木细根表面积的主要分布层,分别有82.4%(侧柏)、86.5%(刺槐)和87.5%(油松)的细根表面积分布。侧柏、刺槐、油松细根表面积垂直分布与剖面土壤水分间呈显著的正相关关系(p0.05)。模型S=AhB(C+Dh+Eh2+Fh3)可以较好地拟合不同树种细根表面积的垂直分布,拟合决定系数R2均在0.85以上。刺槐、侧柏、油松林地土壤含水量的动态变化均表现为10月4月6月8月。刺槐、油松细根表面积在6月出现1个高峰,侧柏在6月和10月各出现1个高峰。树木细根表面积动态与土壤含水量的季节动态不完全一致。侧柏、刺槐、油松生长所需的水分约87%来自降水的补给。但是,总体上侧柏、刺槐、油松细根表面积与林地土壤含水量的相关性不显著(p0.05)。全面了解树木细根季节动态的机理,还需对水分、温度、养分和树种本身遗传特性等影响因子进行综合研究。     

The evolution of root hairs and rhizoids

Background

Almost all land plants develop tip-growing filamentous cells at the interface between the plant and substrate (the soil). Root hairs form on the surface of roots of sporophytes (the multicellular diploid phase of the life cycle) in vascular plants. Rhizoids develop on the free-living gametophytes of vascular and non-vascular plants and on both gametophytes and sporophytes of the extinct rhyniophytes. Extant lycophytes (clubmosses and quillworts) and monilophytes (ferns and horsetails) develop both free-living gametophytes and free-living sporophytes. These gametophytes and sporophytes grow in close contact with the soil and develop rhizoids and root hairs, respectively.

Scope

Here we review the development and function of rhizoids and root hairs in extant groups of land plants. Root hairs are important for the uptake of nutrients with limited mobility in the soil such as phosphate. Rhizoids have a variety of functions including water transport and adhesion to surfaces in some mosses and liverworts.

Conclusions

A similar gene regulatory network controls the development of rhizoids in moss gametophytes and root hairs on the roots of vascular plant sporophytes. It is likely that this gene regulatory network first operated in the gametophyte of the earliest land plants. We propose that later it functioned in sporophytes as the diploid phase evolved a free-living habit and developed an interface with the soil. This transference of gene function from gametophyte to sporophyte could provide a mechanism that, at least in part, explains the increase in morphological diversity of sporophytes that occurred during the radiation of land plants in the Devonian Period.