Response of root respiration and root exudation to alterations in root C supply and demand in wheat

Rising atmospheric CO₂ concentrations have highlighted the importance of being able to understand and predict C fluxes in plant-soil systems. We investigated the responses of the two fluxes contributing to below-ground efflux of plant root-dependent CO₂, root respiration and rhizomicrobial respiration of root exudates. Wheat (Triticum aestivum L., var. Consort) plants were grown in hydroponics at 20°C, pulse-labelled with ¹⁴CO₂ and subjected to two regimes of temperature and light (12 h photoperiod or darkness at either 15°C or 25°C), to alter plant C supply and demand. Root respiration was increased by temperature with a Q ₁₀ of 1.6. Root exudation was, in itself, unaltered by temperature, however, it was reduced when C supply to the roots was reduced and demand for C for respiration was increased by elevated temperature. The rate of exudation responded much more rapidly to the restriction of C input than did respiration and was approximately four times more sensitive to the decline in C supply than respiration. Although temporal responses of exudation and respiration were treatment dependent, at the end of the experimental period (2 days) the relative proportion of C lost by the two processes was conserved despite differences in the magnitude of total root C loss. Approximately 77% of total C and 67% of ¹⁴C lost from roots was accounted for by root respiration. The ratio of exudate specific activity to CO₂ specific activity converged to a common value for all treatments of 2, suggesting that exudates and respired CO₂were not composed of C of the same age. The results suggest that the contributions of root and rhizomicrobial respiration to root-dependent below-ground respiration are conserved and highlight the dangers in estimating short-term respiration and exudation only from measurements of labelled C. The differences in responses over time and in the age of C lost may ultimately prove useful in improving estimates of root and rhizomicrobial respiration.     

Preformation of root primordia in shoots and root morphogenesis in Salix viminalis

Root primordia are formed in the stems of Salix viminalis L. during normal growth. Some of these primordia are produced at definite sites in the nodes. The initiation and early structural and ultrastructural development of the nodal primordia were studied in young shoots. In the fourth node below the terminal leaf cluster some parenchyma cells situated at the lateral leaf gaps formed a small group of initial cells. Derivatives of the newly formed interfascicular cambium added cells to that group, in which later on cell divisions in various directions occurred resulting in the formation of a root primordium. Root morphogenesis was studied in cuttings from one-season-old stems. The cells in the dormant primordia contained many lipid bodies but only a small amount of starch. After the cuttings had been 24 hours in water starch was accumulating in the plastids and lipid bodies were seen in the vacuoles. 48 hours after activation cell divisions occurred throughout the primordia and a layered apical mer-istem was organized. After 96 hours a root cap with amyloplasts was formed and the procambium was well developed. The amyloplasts were sedimented in response to gravity. After six days the first roots were ready to emerge from the stems. Their root caps had a well developed columella and endodermal and pericyclic cells were recognizable.     

Comparative aspects of root and root nodule secondary metabolism in alfalfa

l-phenylalanine ammonia lyase (PAL), indoleacetic acid oxidase (IAA oxidase), O-methyltransferase, peroxidase, total phenolics and total indoles were compared in roots and root nodules of alfalfa. PAL, O-methyl-transferase, total phenolics and total indoles were higher in nodules than in roots. Isolated bacteroids were assayed for O-methyltransferase, PAL, peroxidase and total phenolics, but their levels were either low or not detectable. Nodule leghemoglobin was separated by disc gel electrophoresis and found to have IAA oxidase activity. Phenolics, IAA oxidase and leghemoglobin appear to be interrelated in regulating indole levels in the nodule.     

The course of root differentiation from root primordia in poplar stems

Upon rooting of poplar stem cuttiags the total inductive stimulation does not take place, but the individual root primordia maintain a relative independence. This becomes evident by various time parameters of their differentiation. Under experimental conditions, in dormant cuttings from one year individuals of the hybrid I 214 the activation of the first root primordia occurs after 24 h, the pre-emergent development of the roots formed was completed after 72 h. The activated root primordium is divided into two regions. In the distal region with the predominating cell division the root apex with histogens is formed by the action of initials. From peripheral cell layers of the distal region the so-called “Wurzeltasche” develops which covers the root cap. Due to cell elongation in the proximal region the root apex is pushed up towards the stem surface. The beginning differentiation of the connective vascular tissue is a preparative step for the connection of the vascular system of the developing root with the secondary vascular system of the maternal stem. Following the penetration of the root through the peripheral stem tissues this connection is realized with progressing development. In the developed root the protoxylem elements differentiate continuously and acropetally in direct continuity with tracheids of the basal connective region.     

Earthworm effect on root morphology in a split root system

Plants respond to their environment through adaptations such as root proliferation in nutrient-rich patches. Through their burrows and casts production in soil, earthworms create heterogeneity which could lead to local root adaptations or systemic effects. To investigate the effect of earthworms on root system morphology and determine whether earthworm effect is local or systemic, we set up two independent split root experiments with rice or barley, (i) without earthworm (CC), (ii) with earthworms in both compartments (EE), and (iii) with earthworms in one single compartment (CE). Earthworms had an effect on belowground plant biomass. The relative length of thick roots decreased with an increasing abundance of earthworms. Some root diameter classes responded to earthworm number in a linear or curvilinear way, making simple conclusions difficult. We found no difference in root biomass or morphology between the two compartments of the split root system in the CE treatment, but a positive effect of earthworm biomass on root biomass, volume, surface area, and length at the whole plant level. Results supported a systemic effect dependent on earthworm abundance. Modification of nutrient mineralization, soil physical structure, and/or the concentration of signal molecules could all be responsible for this systemic effect.     

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.     

Comparative anatomy of root meristem and root cap in some species of Podostemaceae and the evolution of root dorsiventrality

In the unusual aquatic Podostemaceae, the root is the leading organ of the plant body and is variously compressed and submerged as it adheres to rock surfaces in rapid water. In an anatomical comparison of the root apical meristems and root caps of 33 species that represent the major lineages of the family, the dorsiventrality of root meristems varied and was classified into four patterns: (1) The root cap is produced outward from a nearly radially symmetrical meristem. (2) The meristem and root cap are markedly dorsiventral; the outermost cells of the hood-shaped cap are acroscopic derivatives from bifacial initials on the ventral side, while the pattern on the dorsal side is similar to pattern 1. (3) Bifacial initials are on both the dorsal and ventral sides. (4) No root cap is present. An evolutionary polarity may be evident from pattern 1 to 2 and then to 3. Pattern 2 arose in the early evolution of the subfamily Podostemoideae and subsequently, pattern 3 arose in species with crustose roots, while the least specialized pattern 1 is retained in Tristichoideae and Weddellinoideae. Pattern 4 characterized by caplessness may have appeared recurrently in Tristichoideae and Podostemoideae. These evolutionary changes in the meristem preceded the specialization of external root morphologies.     

根修剪对黄土旱塬冬小麦(Triticum aestivum)根系分布、根系效率及产量形成的影响

通过田间试验研究了不同时期根修剪处理对冬小麦（Triticum aestivum）根系大小与分布、根系效率、水分利用效率及产量形成的影响。设置4个根修剪处理：越冬期小剪根（WS）、越冬期大剪根（WB）,返青期小剪根（GS）、返青期大剪根（GB）,未剪根小麦作为对照（CK）。结果表明,到花期时,各根修剪处理小麦的在0～120cm总根量均显著小于对照。与对照相比各根修剪处理主要是显著地减少了上层土壤中的根量。但WS和GS两小剪根处理和对照相比在中层土壤中有较大的根量;花后各处理小麦旗叶的气孔导度和蒸腾速率均显著大于对照。这说明根修剪处理减少了小麦表层的根量,从而削弱了表土干旱信号对作物与外界气体交换的抑制作用。花期时各根修剪小麦的净光合速率均显著高于对照,而单位面积上的根呼吸速率均显著小于对照,根修剪处理提高了小麦的根系效率,使更多的光合产物用于籽粒生产,从而提高了小麦的收获指数。根修剪还提高了小麦的水分利用效率,其中WS、WB、GS处理的水分利用效率显著高于对照。但是GB处理的水分利用效率却没有显著提高。因此,本研究进一步证明了由不同年代品种得到的推测,认为在旱地农业中,通过遗传育种或采用适当农艺措施优化根系分布,既可以减少生长前期作物对水分的过度消耗,又能够削弱花后表土过度干旱对作物生长抑制作用,同时降低根系对同化产物的消耗,对作物产量及水分利用效率的提高具有积极的作用。     

施肥对日本落叶松人工林细根直径、根长和比根长的影响

以辽宁东部山区16年生日本落叶松人工林为研究对象,探讨施肥对日本落叶松1～5级根序中细根直径、根长和比根长的影响.结果表明：随着根序等级的增加,日本落叶松细根平均直径和根长显著增加（P<0。05,P<0。01）、比根长则显著下降（P<0。01）.在日本落叶松的5级根序中,1级根的平均直径最细、根长最短、比根长最高,而5级根则相反;随着根序等级的增加,日本落叶松细根平均直径、根长和比根长的变异系数逐渐增大.除1级根外,土层对细根的平均直径、根长和比根长没有显著影响(P>0。05).与对照样地相比,施肥对各级细根平均直径、根长和比根长的影响主要表现在1～2级根上,对3级根序以上的细根影响不显著（P>0.05）.其中,施氮肥显著降低了1～2级根的平均直径（P<0.05）,施氮肥以及氮磷肥显著降低了表层土壤（0～10 cm）中1级根的平均根长（P<0.05）,表层土壤中细根的比根长在施氮肥的条件下显著增加（P<0.05）.     

Role of cytokinin and auxin in shaping root architecture: regulating vascular differentiation, lateral root initiation, root apical dominance and root gravitropism

BACKGROUND AND AIMS: Development and architecture of plant roots are regulated by phytohormones. Cytokinin (CK), synthesized in the root cap, promotes cytokinesis, vascular cambium sensitivity, vascular differentiation and root apical dominance. Auxin (indole-3-acetic acid, IAA), produced in young shoot organs, promotes root development and induces vascular differentiation. Both IAA and CK regulate root gravitropism. The aims of this study were to analyse the hormonal mechanisms that induce the root's primary vascular system, explain how differentiating-protoxylem vessels promote lateral root initiation, propose the concept of CK-dependent root apical dominance, and visualize the CK and IAA regulation of root gravitropiosm. KEY ISSUES: The hormonal analysis and proposed mechanisms yield new insights and extend previous concepts: how the radial pattern of the root protoxylem vs. protophloem strands is induced by alternating polar streams of high IAA vs. low IAA concentrations, respectively; how differentiating-protoxylem vessel elements stimulate lateral root initiation by auxin-ethylene-auxin signalling; and how root apical dominance is regulated by the root-cap-synthesized CK, which gives priority to the primary root in competition with its own lateral roots. CONCLUSIONS: CK and IAA are key hormones that regulate root development, its vascular differentiation and root gravitropism; these two hormones, together with ethylene, regulate lateral root initiation.     

An unusual root tip formation in hairy root culture of Hyoscyamus muticus

Summary Hairy root cultures of Hyoscyamus muticus were established using Agrobacterium rhizogenes ATCC 15834. In one out of 8 clones established, an unusual root tip formation was observed after transfer of cultures from half-strength Murashige and Skoog (1962) to White's medium (1939). This phenomenon was associated with the production of a fine brownish cell suspension culture. Hairy root development resumed after transfer of the root tips from White to half-strength Murashige and Skoog medium. After plating the isolated brownish cells on hormone-free half-strength Murashige and Skoog or White solid medium, callus proliferation was observed, and then redifferentiation of hairy roots occurred. The polymerase chain reaction analysis of the H. muticus hairy root (clone Z2) revealed that only the tl region of the T-DNA was integrated. The growth and the production of five tropane alkaloids by this clone were examined.Abbreviations PCR Polymerase Chain Reaction - MS medium Murashige and Skoog Medium - 1/2 MS medium half-strength MS medium - WP medium Woody Plant medium - RC medium Root Culture medium - WH medium White medium - HPLC High Performance Liquid Chromatography - wt. weight     

Hydrogenase in actinorhizal root nodules and root nodule homogenates.

Hydrogenases were measured in intact actinorhizal root nodules and from disrupted nodules of Alnus glutinosa, Alnus rhombifolia, Alnus rubra, and Myrica pensylvanica. Whole nodules took up H2 in an O2-dependent reaction. Endophyte preparations oxidized H2 through the oxyhydrogen reaction, but rates were enhanced when hydrogen uptake was coupled to artificial electron acceptors. Oxygen inhibited artifical acceptor-dependent H2 uptake. The hydrogenase system from M. pensylvanica had a different pattern of coupling to various electron acceptors than the hydrogenase systems from the alders; only the bayberry system evolved H2 from reduced viologen dyes.     

Programmed cell death in the root cortex of soybean root necrosis mutants

The soybean root necrosis (rn) mutation causes a progressive browning of the root soon after germination that is associated with accumulation of phytoalexins and pathogenesis-related proteins and an increased tolerance to root-borne infection by the fungal pathogen, Phytophthora sojae. Grafting and decapitation experiments indicate that the rn phenotype is root-autonomous at the macroscopic level. However, the onset and severity of browning was modulated in intact plants by exposure to light, as was the extent of lateral root formation, suggesting that both lateral roots and the rn phenotype could be directly or indirectly controlled by similar shoot-derived factors. Browning first occurs in differentiated inner cortical cells adjacent to the stele and is preceded by a wave of autofluorescence that emanates from cortical cells opposite the xylem poles and spreads across the cortex. Before any visible changes in autofluorescence or browning, fragmented DNA was detected by TUNEL (T erminal deoxynucleotidyl transferase-mediated dU TP-digoxigenin n ick e nd l abeling) in small clusters of inner cortical cells that subsequently could be distinguished cytologically from neighboring cells throughout rn root development. Inner cortical cells overlying lateral root primordia in either Rn or rn plants also were stained by TUNEL. Features commonly observed in animal cell apoptosis were confirmed by electron microscopy but, surprisingly, cells with a necrotic morphology were detected alongside apoptotic cells in the cortex of rn roots when TUNEL-positive cells were first observed. The two morphologies may represent different stages of a common pathway for programmed cell death (pcd) in plant roots, or two separate pathways of pcd could be involved. The phenotype of rn plants suggests that the Rn gene could either negatively regulate cortical cell death or be required for cortical cell survival. The possibility of a mechanistic link between cortical cell death in rn plants and during lateral root emergence is discussed.     

Effects of root diameter and root nitrogen concentration on in situ root respiration among different seasons and tree species

Knowledge of root respiration is a prerequisite for a better understanding of ecosystem carbon budget and carbon allocation. However, there are not many relevant data in the literature on direct measurements of in situ root respiration by root chamber method. Furthermore, few studies have been focused on the effects of root diameter (D _r) and root nitrogen concentration (N _r) on in situ root respiration among different seasons and tree species. To address these goals, we used a simplified root-chamber system to measure in situ root respiration rates of Acacia crassicarpa and Eucalyptus urophylla in subtropical plantations of south China. We found that the species and season variation in root respiration were affected by D _r and N _r. Also, the root respiration per unit dry mass (R _r, nmol CO₂ g⁻¹ s⁻¹) and root respiration per unit N (R _n, nmol CO₂ g N⁻¹ s⁻¹) were affected by D _r and N _r. The R _r, R _n, N _r and soil temperature sensitivity (Q ₁₀) of R _r for the two species significantly decreased with an increase of D _r. The R _r of the two species showed significant an inter-seasonal and diurnal pattern, and this trend decreased with increasing D _r. Both the R _r and Q ₁₀ of the two species increased with increasing N _r. The D _r and N _r explained 54 and 52% of the observed variation in R _r for A. crassicarpa, and 65 and 70% for E. urophylla. The R _r, N _r, and Q ₁₀ of A. crassicarpa were significantly higher than those of E. urophylla. Our results indicated that root respiration was dependent on D _r and N _r, and this dependence varied with season and plant species.     

Association of soybean root mycoflora with root modulation, root necrosis and plant fresh weight

Fungi were isolated from soybean plants in ten fields in south-west France. Correspondence analysis (COA) was used to find associations among taxa (species, genera, groups) and sampling units (i.e. one field at one sampling date). The first axis of the COA demonstrated a dominant sampling date effect; the second indicated an association among certain sampling units and specific taxa. Regression analysis of COA coordinate values indicated that fields with high relative isolation frequencies of Fusarium spp., F. oxysporum and ‘other species’ (mostly Chaetomium spp. and sterile fungi) were characterised by greater plant fresh weight and nodule number and lower ratings for root necrosis. Cylindrocarpon spp., Cephalosporium spp. and F. solanihad high relative isolation frequencies in fields with lower plant weight, lower nodule number and higher root necrosis ratings. The relative importance of fungal taxa was also related to soil type.     

Timing of root dormancy and tolerance to root waterlogging in clonal Sitka spruce

 Actively growing root tips of Picea sitchensis (Bong.) Carr. plants are highly susceptible to damage if waterlogged, but they are known to have some tolerance after they stop growing in the autumn. This paper describes the selection of clones on the basis of root dormancy timing and the corresponding responses of their roots to over-winter waterlogging. Sitka spruce transplants of Alaska, Queen Charlotte Islands (QCI), and Washington provenances were screened for early or late root dormancy over 2 successive years. Cuttings were propagated from the selected plants and after growing on for 2 years, they were planted in transparent acrylic tubes within outdoor ‘root observation chambers’. Extension of main roots and the timing of onset of root dormancy was recorded on the clonal plants. The tubes were flooded in November and maintained with a water table 280 mm below the soil surface until March of the next year. Waterlogging caused most main root tips to die back, but within 2 months of draining regeneration occurred on the main roots below the waterlogging level. This regeneration was most commonly the growth of existing lateral tips or production of new lateral roots. Roots of early-dormant Washington plants died back on average 129 mm less than late-dormant Washington plants, and early-dormant Alaska plants had 173 mm less dieback than late-dormant Alaska plants. Differences between the clones of the QCI provenance were not significant. The 40% and 52% increases in survival depth of roots in early-dormant Washington and Alaska clones respectively indicates a potential for improving the rooting depth of Sitka spruce on seasonally waterlogged soils by planting clones selected on the basis of root dormancy. Received: 14 July 1997 / Accepted: 15 September 1997     

水曲柳根系生物量、比根长和根长密度的分布格局

采用连续钻取土芯法在生长季内对东北林业大学帽儿山实验林场17年生水曲柳人工林根系取样,研究水曲柳不同直径根系现存生物量、比根长和根长密度及垂直分布状况.结果表明,水曲柳人工林根系总生物量为1 637.6 g·m^-2,其中活根生物量占85%,死根占15%.在活根生物量当中,粗根(直径5～30 mm)占的比例最高（69.95%）,其次为活细根（直径<1 mm,13.53%）,小根(1～2 mm)和中等直径的根(2～5 mm)比例较小（分别为7.21%和9.31%）.直径<1 mm活细根的比根长为32.20 m·g^-1,直径5～30 mm粗根的比根长为0.08 m·g^-1.单位面积上活根的总长度为6 602.54 m·m^-2,其中直径<1 mm的细根占92.43%,其它直径等级则不到活根总长度的8%.直径<1 mm的细根生物量与根长密度具显著线性关系（R²=0.923）,但与比根长无显著相关关系（R²=0.134）.     

Photosynthesis and root growth in Spartina alterniflora in relation to root zone aeration

Spartina alterniflora Lois. is a dominant species growing in intermediate and saline marshes of the US Gulf coast and Atlantic coastal marshes. S. alterniflora plants were subjected to a range of soil redox potential (Eh) conditions representing a well aerated to reduced conditions in a rhizotron system under controlled environmental conditions. The low soil Eh resulted in inhibition of root elongation shortly after treatment initiation. Root elongation was reduced as soil Eh approached values below ca. +350 mV. Substantial decrease in root elongation was noted when soil Eh fell below +200 mV. Generally, net photosynthetic rate (PN) decreased as soil Eh was reduced, with substantial reductions in PN found when Eh approached negative values. Average PN was reduced to 87, 64, and 44% of control under +340, +245, and -180 mV treatments, respectively. The reductions in root elongation and PN in response to low soil Eh indicated the adverse effects of low soil Eh on plant functioning and the need for periods of soil aeration that allow plants to resume normal functioning. Thus periods of drainage allowing soil aeration during the growing season appear to be critical to S. alterniflora by providing favorable conditions for root growth and gas exchange with important implications for plant carbon fixation. This revised version was published online in June 2006 with corrections to the Cover Date.