A Comparison of Dry Matter Partitioning in Seminal and Adventitious Plants of Trifolium repens L

Trifolium repens has two types of root, one derived from theseed and the adventitious roots derived from the stolon nodes.It has been suggested that these two systems have differentpotentials for supporting growth. This paper presents a comparisonof plants grown on single seedling or adventitious roots anddemonstrates that although the shoot: root ratios for the twotypes differ this may be explained by differing shoot morphologies.Comparison of the lamina: root ratios for the two types of plantproduced no statistically significant differences and it isproposed that the two types of root system do not differ intheir relationship with leaf growth. A mechanism for large diameter‘tap’ root formation is suggested. white clover, Trifolium repens L., adventitious roots, seedling roots, shoot: root ratio     

Effects of sand burial depth on the root system of Salix cheilophila seedlings in Mu Us Sandy Land, Inner Mongolia, China

Salix cheilophila Schneid. is a naturally occurring Salix species in Mu Us Sandy Land, Inner Mongolia, China. We focused on the morphological adaptability of S. cheilophila to sand dune burial. For morphological measurements, 32 S. cheilophila seedlings were removed from a community which was in the process of being buried by a shifting sand dune. Each seedling collected included the entire root system. We measured the number, length, and biomass of the adventitious roots, primary lateral roots, and taproot, and compared the morphological characteristics of the root system, including adventitious roots, for seedlings buried to various levels in the sand. The growth range of adventitious roots increased as the length of the buried portion of the main shoot increased. In addition, the total dry weight of all current-year shoots tended to increase gradually with increasing total dry weight of the adventitious roots. These results suggest that S. cheilophila tends to make use of the sedimentary sand layer that accompanies shifting sand dunes. However, there was no correlation between biomass or number of adventitious roots and the length of the buried part of the main shoot. Thus, S. cheilophila does not grow adventitious roots proportional to the buried part. These morphological characteristics of the root system, including the adventitious roots, may indicate that S. cheilophila has poor morphological adaptability to sand dune burial.     

Contribution of adventitious vs initial roots to growth and physiology of black spruce seedlings

Black spruce (Picea mariana [Mill.] BSP) is a boreal tree species characterized by the formation of an adventitious root system. Unlike initial roots from seed germination, adventitious roots gradually appear above the root collar, until they constitute most of mature black spruce root system. Little is known about the physiological role they play and their influence on tree growth relative to initial roots. We hypothesized that adventitious roots present an advantage over initial roots in acquiring water and nutrients. To test this hypothesis, the absorptive capacities of the two root systems were explored in a controlled environment during one growing season. Black spruce seedlings were placed in a double‐pot system allowing irrigation (25 and 100% water container capacity) and fertilization (with or without fertilizer) inputs independent to initial and adventitious roots. After 14 weeks, growth parameters (height, diameter, biomass), physiology (net photosynthetic rate, stomatal conductance, shoot water potential) and nutrient content (N, P, K, Ca and Mg foliar content) were compared. Most measured parameters showed no difference for the same treatment on adventitious or initial roots, except for root biomass. Indeed, fertilized black spruce seedlings invested heavily in adventitious root production, twice as much as initial roots. This was also the case when adventitious roots alone were irrigated, while seedlings with adventitious roots subjected to low irrigation produced initial root biomass equivalent to that of adventitious roots. We conclude that black spruce seedlings perform equally well through adventitious and initial roots, but if resources are abundant, they strongly promote development of adventitious roots.     

冰凉花根系发育形态学的初步研究

冰凉花属于宿根植物,其根几乎终生为初生结构。一年生苗直根系的主根和侧根的初生木质部均为二原型。二至多年生植株的不定根分为二原型、三原型、四原型和五原型,其分枝均为二原型。根在一年中有两个生长期。展叶结果营养期产生新的不定根,旧根顶端恢复延长生长或产生新的分枝。夏季枯萎休眠期新生根变成黄褐色,停止生长。秋季地下生长期根系又开始生长。冬季严寒迫使生长趋于停止。研究冰凉花根系发育形态学不仅具有理论意义而且有很好的应用前景。     

Compensatory function for water transport by adventitious roots of <Emphasis Type="Italic">Ipomoea pes-caprae</Emphasis>

To determine the role of adventitious roots in supplying water to Ipomoea pes-caprae (L.) Sweet (Convolvulaceae), we examined the effects of water deficit on water uptake and the growth patterns of leaves and shoots. After stopping the water supply from the primary root or adventitious roots, the water-uptake rate of the other root system increased steeply within 90–100 min to a level of 90% of the pretreatment water-uptake rate of the whole plant. Thus, the primary and adventitious roots can compensate for a decrease in the water-uptake rate of the whole plant caused by dehydration. The continuous growth of leaves and shoots after dehydration suggests that an increase in the water-uptake rate by either root system can support plant growth, although the growth rates of immature leaves in plants with no water supply from the primary or adventitious roots were lower than in controls. We conclude that the water supply from adventitious roots contributes to the survival and growth of plants, and will be important for vegetative propagation.     

Influence of galactoglucomannan oligosaccharides on root culture of <Emphasis Type="Italic">Karwinskia humboldtiana</Emphasis>

Galactoglucomannan oligosaccharides (GGMOs) activity in K. humboldtiana root culture has been determined. GGMOs inhibited adventitious root growth and lateral root induction in contrast to IAA, IBA, and NAA stimulating effect in these processes. Similarly, the combination of GGMOs with natural auxins (IAA, IBA) evoked an inhibition of adventitious root growth and lateral root induction that depended on the oligosaccharides concentration and the type of auxin. The growth stimulating effect of the synthetic auxin, NAA, in adventitious roots was negatively affected by GGMOs, but they were without influence on lateral root induction. The presence of oligosaccharides triggered lateral root position on adventitious roots and the anatomy of adventitious roots (diameter, proportion of primary cortex to the central cylinder, number and size of primary cortical cells, intercellular spaces, and the number of starch grains in cells of primary cortex) in dependence on their coactions with auxin.     

Growth responses to waterlogging and drainage of woody Hakea (Proteaceae) seedlings,originating from contrasting habitats in south-western Australia

We investigated the responses of seven woody Hakea (Proteaceae) species (two populations of each), to two months of waterlogging and subsequent drainage, in a controlled glasshouse experiment. The species originated from contrasting environments (winter-wetland versus non-wetland habitats), and differed in abundance (endangered ironstone species versus common species). Waterlogging arrested growth of the main root system, and stimulated the formation of superficial adventitious roots just below the root/shoot junction in all species. Wetland species produced at least twice the amount of adventitious root dry mass of that of non-wetland species, due to differences in number, length or degree of branching. Their adventitious roots also tended to have higher porosities (7–10% versus 5–6% gas spaces). The relative amount of adventitious roots formed was strongly, positively correlated with the maintenance of shoot growth, and only the non-wetland species showed significant shoot growth reductions (19%) upon waterlogging. Dry mass percentage of stems and leaves, and leaf dry mass per area (LMA) increased considerably during waterlogging in all species (averages of 15, 29 and 27%, respectively), returning to the values of continuously drained control plants after drainage. Similarly, upon drainage, a suppression of shoot growth (average 35%) and a stimulation of root growth (average 50%) restored the root mass ratios to those of control plants. We found a negative correlation between the maintenance of growth during waterlogging versus that after waterlogging, suggesting a trade-off in functioning of the superficial adventitious roots between waterlogged and drained conditions. The rare winter-wet ironstone endemics resembled the common winter-wet species in most of their responses to waterlogging and drainage. Therefore, the results presented here cannot offer an explanation for their different distribution patterns. Our results suggest that non-wetland species may be disadvantaged in a wetland environment, due to their lower capacity to form adventitious roots resulting in stronger growth reductions.     

Aquatic adventitious root development in partially and completely submerged wetland plants Cotula coronopifolia and Meionectes brownii

Background and Aims

A common response of wetland plants to flooding is the formation of aquatic adventitious roots. Observations of aquatic root growth are widespread; however, controlled studies of aquatic roots of terrestrial herbaceous species are scarce. Submergence tolerance and aquatic root growth and physiology were evaluated in two herbaceous, perennial wetland species Cotula coronopifolia and Meionectes brownii.

Methods

Plants were raised in large pots with ‘sediment’ roots in nutrient solution and then placed into individual tanks and shoots were left in air or submerged (completely or partially). The effects on growth of aquatic root removal, and of light availability to submerged plant organs, were evaluated. Responses of aquatic root porosity, chlorophyll and underwater photosynthesis, were studied.

Key Results

Both species tolerated 4 weeks of complete or partial submergence. Extensive, photosynthetically active, aquatic adventitious roots grew from submerged stems and contributed up to 90 % of the total root dry mass. When aquatic roots were pruned, completely submerged plants grew less and had lower stem and leaf chlorophyll a, as compared with controls with intact roots. Roots exposed to the lowest PAR (daily mean 4·7 ± 2·4 µmol m⁻² s⁻¹) under water contained less chlorophyll, but there was no difference in aquatic root biomass after 4 weeks, regardless of light availability in the water column (high PAR was available to all emergent shoots).

Conclusions

Both M. brownii and C. coronopifolia responded to submergence with growth of aquatic adventitious roots, which essentially replaced the existing sediment root system. These aquatic roots contained chlorophyll and were photosynthetically active. Removal of aquatic roots had negative effects on plant growth during partial and complete submergence.     

入侵植物水花生解剖学和组织化学染色研究

水花生(Alternanthera philoxeroides)因其表型可塑性、高生长速率和快速无性繁殖能适应水、陆生境.该文利用光学显微镜和荧光显微镜对水、陆生境的水花生不定根、茎解剖结构、组织化学特征及质外体通透性进行了研究.结果表明:(1)水生境下,其不定根皮层中具较大裂生型通气组织,无次生生长,内皮层具凯氏带且...     

ANATOMY OF SEEDLING ROOTS OF PSEUDOTSUGA MENZIESII

The seedling root system of Pseudotsuga menziesii (Mirb.) Franco consists of the primary root, active long laterals, long laterals that become mycorrhizal, and short roots that may or may not become mycorrhizal. Numerous adventitious roots arise from the pericycle in young roots and from the vascular cambium and pericycle in older roots following pruning. All actively growing apices have a single plate of initials, a complex zonation of mother cells, and a similar pattern of primary tissue differentiation. Short roots and mycorrhizal short roots have 2 plates of initials, one producing the stele and the other the root cap and cortex, and differentiation occurs close to the apex. Primary and adventitious roots are usually triarch, while long laterals are usually diarch as are all short roots. The latter lack secondary xylem, but mycorrhizal short roots may produce a small amount of secondary phloem.     

Seedling developmental anatomy of an undescribed Malaccotristicha species (Podostemaceae, subfamily Tristichoideae) with implications for body plan evolution

The seedling development of an undescribed Malaccotristicha species was observed by using seedling culture and microtomy to infer the evolution of body plan with a focus on the root, which is a developmentally leading organ of most Podostemaceae. The young seedling has a small primary shoot apical meristem and a primary root apical meristem. The shoot meristem develops into a plumular ramulus, and the root meristem, into a cylindrical radicle with no root cap. The radicle transforms to a dorsiventral, flattened, capped primary root. An adventitious root develops endogenously on the lateral side of the hypocotyl and is similar to the primary root. This is a new pattern in Podostemaceae. Comparison of this and described patterns of Podostemaceae (and the sister-group Hypericaceae) suggests that the radicle was lost in the early evolution of Podostemaceae and instead adventitious roots replaced it as a leading organ.     

Cytokinin receptors are required for normal development of auxin-transporting vascular tissues in the hypocotyl but not in adventitious roots

Plants alter the architecture of their root systems to adapt to the environment by modulating post-embryonic (lateral and adventitious) root formation and growth. To understand better the genetic basis of this regulation, we screened ethylmethane sulfonate-mutagenized lines of Arabidopsis thaliana for adventitious rooting mutants. One mutant showed retardation of the primary root growth, no production of lateral roots and enhanced formation of adventitious roots. Mapping and genetic complementation revealed that this mutant named wooden leg-3 (wol-3) was an allele of ARABIDOPSIS HISTIDINE KINASE 4 (AHK4), a locus known to encode a cytokinin receptor. Although the vascular system of the primary root and hypocotyl in the wol-3 mutant was aborted, that of the adventitious roots was normally developed. In the hypocotyl of the wol-3 mutant, auxin signals accumulated around the aborted vascular system. The application of auxin to primary roots induced lateral root formation in the wol-3 mutant. Transport of radiolabeled auxin from the top of the hypocotyl to the primary root was inhibited in wol-3. Although only a single amino acid alteration had occurred in AHK4, the root morphology in the wol-3 mutant was quite similar to that in the ahk2 ahk3 ahk4 triple mutant, which is a loss-of-function mutant of the three cytokinin receptors. This implies that the functional disturbance of AHK4 affects the function of the other receptors. Our results suggest that cytokinin receptors are necessary for the formation of auxin-transporting vascular tissues in the hypocotyl, but not in adventitious roots.     

Optimization and quality assessment of adventitious roots culture in Panax quinquefolium L.

In this study, adventitious roots of Panax quinquefolium L. have been successfully established. The highest induction rate of roots was obtained in MS medium containing 3 mg L^?1 IBA after 4 weeks of culture. The culture conditions of adventitious root were optimized and evaluated with response surface methodology. The best culture conditions for root growth seemed to be 0.75 salt strength MS medium, 4.70 g L^?1 inoculum size and 40 days of culture. The active component contents of adventitious roots were compared with those of native roots. The total saponins content was found to be 16.28 mg g^?1 in native root and 4.64 mg g^?1 in adventitious root. The polysaccharide content of the adventitious root was 1.5 times higher than that in the native P. quinquefolium (30.54 vs. 20.28 mg g^?1).     

Dependency of Iron Reduction on Development of a Unique Root Morphology in Ficus benjamina L

The activity of the Fe³⁺ reductase of excised adventitious roots of Ficus benjamina L., grown in hydroponic culture without iron, was determined by a colorometric assay simplified by the use of a microplate reader. Reductase activity remained the same from pH 4.5 to 6.5 and decreased sharply above pH 6.5. Acetate buffer inhibited reduction. During early stages of root growth, excised roots did not exhibit Fe³⁺ reductase activity. After several weeks and extensive root system development, Fe³⁺ reduction still was not detectable in primary roots, but intermediate and high rates of reduction occurred in lateral and newly formed root clusters, respectively. Clustered roots only developed on plants grown at 0 or very low (<1 micromolar) iron. Microscopic examination revealed the root cluster to be composed of up to 30 lateral roots, usually less than 1 millimeter in diameter and 1 centimeter in length, that were completely covered with root hairs.     

Seedling growth variation in response to sand burial in four <Emphasis Type="Italic">Artemisia</Emphasis> species from different habitats in the semi-arid dune field

We compared seedling growth of four Artemisia species dominated at different habitats to determine whether interspecific seedling growth variation of a same genus in tolerance to burial can be used to explain plant distribution in the sand dune field. Interdune lowland species, Artemisia gmelinii, stabilized dune species, A. frigida, semi-stabilized dune species, A. halodendron, and active dune species, A. wudanica were selected. Seedlings grown for 3 weeks were treated at five burial depths for three burial times in pot experiments. Species from the habitats with little burial had smaller survival rate, dry weight and stem elongation speed than those from the habitats with intensive burial when buried. Furthermore, when buried, the former tended to adjust biomass allocation between shoot and root and produce adventitious buds, while the latter tended to maintain a constant root:shoot ratio and produce adventitious roots. We conclude that (1) seedlings of species with a long evolutionary history of exposure to sand burial (from the active sand dune), show quicker stem growth when buried than do seedlings of species from the habitats with little or no sand burial; (2) seedlings of species which do not change root:shoot ratio might be more tolerant of sand burial than those do; (3) seedlings of species from the habitats with intensive sand burial is prone to produce adventitious roots and seedlings of species from the habitats with little or no sand burial tend to produce adventitious buds when buried.     

Relationships between the development of adventitious roots and the biosynthesis of anthocyanins in first internodes of sorghum

The initiation and subsequent growth of adventitious roots in excised first internodes of Sorghum vulgare var. Wheatland milo were studied to determine the effect of these processes on anthocyanin biosyntheses. Segmentation of the internodes inhibited both adventitious root growth and accumulation of cyanidin equally in all segments; these results can be interpreted as a common requirement for bidirectional longitudinal transport. The presence of the coleoptile, especially in the absence of the base of the internode, inhibited the growth of the roots, but increased the number of root initials. High intensities of white and blue light which induced cyanidin synthesis slightly decreased adventitious root growth. Anaerobic conditions produced by solution infiltration strongly inhibited the growth of adventitious roots and greatly increased the accumulation of apigeninidin and luteolinidin. Addition of indoleacetic acid, kinetin and cofactors such as pyridoxine produced effects on the initiation and subsequent growth of these roots similar to those effects reported in the literature. But unlike root formation in hypocotyls, the initiation of adventitious roots in Sorghum internodes was not always directly correlated with the accumulation of anthocyanins, and the subsequent growth of these roots was frequently inversely correlated with some of the anthocyanin biosyntheses. The possible nature of these correlations is discussed. Comparisons are made with related Sorghum lines and mutants.     

Gibberellin disturbs the balance of endogenesis hormones and inhibits adventitious root development of Pseudostellaria heterophylla through regulating gene expression related to hormone synthesis

The adventitious roots of some plants will develop into tuberous roots which are widely used in many traditional Chinese medicines, including Pseudostellaria heterophylla. If adventitious root development is inhibited, the yield of Chinese medicinal materials will be reduced. Gibberellic acid is an important phytohormone that promotes plant growth and increases the resistance to drought, flood or disease. However, the effects of gibberellic acid on adventitious roots of Pseudostellaria heterophylla are not clear. Here, we reports GA3 suppressed adventitious root development of Pseudostellaria heterophylla by disturbing the balance of endogenesis hormones. By detecting the contents of various endogenous hormones, we found that the development of adventitious roots negatively correlated with the content of CA3 in tuberous roots. Exogenous GA3 treatment decreased the diameter of adventitious roots, but increased the length of adventitious roots of Pseudostellaria heterophylla. In contrast, blocking the biosynthesis of GA3 suppressed stem growth and promoted the xylem of tuberous roots development. Moreover, exogenous GA3 treatment resulted in imbalance of endogenesis hormones by regulating their synthesis-related genes expression in xylem of tuberous roots. These results suggest GA3 broke the established distribution of hormones by regulating synthesis, transport and biological activation of hormones to activate the apical meristem and suppress lateral meristem. Regulating GA3 signaling during adventitious roots development would be one of the possible ways to increase the yield of P. heterophylla.     

Comparative developmental anatomy of seedlings in nine species of podostemaceae (subfamily Podostemoideae)

The developmental anatomy is described for seedlings of nine Asian and Australian species of Podostemaceae, subfamily Podostemoideae. The hypocotyl is rudimentary (except in Zeylanidium olivaceum) and does not form a primary root in any of the species examined. An adventitious root forms endogenously in the hypocotyl of six species with ribbon-like or flattened subcylindrical roots, and in Z olivaceum with foliose roots. In contrast, it forms exogenously in Hydrobryum griffithii and Synstylis micranthera with foliose roots. The juvenile root becomes flattened and dorsiventral, branches exogenously (in Polypleurum stylosum, P. wallichii and Z. lichenoides) and produces shoots endogenously (in P. stylosum, P. wallichii, S. micranthera and Z. lichenoides). The root meristem is simple, composed of surface and uniform inner cells, and is devoid of root cap initials in all species. The reduced meristem morphology of seedling roots may be primitive in the Asian-Australian Podostemoideae. A root cap or protective tissue did not form during the culture period, even in the seven species with capped adult roots, probably due to its delayed development. It was absent throughout ontogeny in the other two species. No obvious shoot apical meristem forms between the cotyledons. One to several leaves occupy the shoot apical area in species with endogenous adventitious roots, while no leaves are formed in species with exogenous roots. These differences suggest recurrent origins of foliose roots in the Asian clade. Similarities between the unique seedling morphology and mutant Arabidopsis phenotypes are discussed.     

Architectural Tradeoffs between Adventitious and Basal Roots for Phosphorus Acquisition

Adventitious rooting contributes to efficient phosphorus acquisition by enhancing topsoil foraging. However, metabolic investment in adventitious roots may retard the development of other root classes such as basal roots, which are also important for phosphorus acquisition. In this study we quantitatively assessed the potential effects of adventitious rooting on basal root growth and whole plant phosphorus acquisition in young bean plants. The geometric simulation model SimRoot was used to dynamically model root systems with varying architecture and C availability growing for 21 days at 3 planting depths in 3 soil types with contrasting nutrient mobility. Simulated root architectures, tradeoffs between adventitious and basal root growth, and phosphorus acquisition were validated with empirical measurements. Phosphorus acquisition and phosphorus acquisition efficiency (defined as mol phosphorus acquired per mol C allocated to roots) were estimated for plants growing in soil in which phosphorus availability was uniform with depth or was greatest in the topsoil, as occurs in most natural soils. Phosphorus acquisition and acquisition efficiency increased with increasing allocation to adventitious roots in stratified soil, due to increased phosphorus depletion of surface soil. In uniform soil, increased adventitious rooting decreased phosphorus acquisition by reducing the growth of lateral roots arising from the tap root and basal roots. The benefit of adventitious roots for phosphorus acquisition was dependent on the specific respiration rate of adventitious roots as well as on whether overall C allocation to root growth was increased, as occurs in plants under phosphorus stress, or was lower, as observed in unstressed plants. In stratified soil, adventitious rooting reduced the growth of tap and basal lateral roots, yet phosphorus acquisition increased by up to 10% when total C allocation to roots was high and adventitious root respiration was similar to that in basal roots. With C allocation to roots decreased by 38%, adventitious roots still increased phosphorus acquisition by 5%. Allocation to adventitious roots enhanced phosphorus acquisition and efficiency as long as the specific respiration of adventitious roots was similar to that of basal roots and less than twice that of tap roots. When adventitious roots were assigned greater specific respiration rates, increased adventitious rooting reduced phosphorus acquisition and efficiency by diverting carbohydrate from other root types. Varying the phosphorus diffusion coefficient to reflect varying mobilities in different soil types had little effect on the value of adventitious rooting for phosphorus acquisition. Adventitious roots benefited plants regardless of basal root growth angle. Seed planting depth only affected phosphorus uptake and efficiency when seed was planted below the high phosphorus surface stratum. Our results confirm the importance of root respiration in nutrient foraging strategies, and demonstrate functional tradeoffs among distinct components of the root system. These results will be useful in developing ideotypes for more nutrient efficient crops.     

ORIGIN AND DEVELOPMENT OF LATERAL ROOTS IN TYPHA GLAUCA

Lateral roots of Typha glauca arose from the pericycle of the parent adventitious root. Periclinal divisions of the pericycle gave rise to two layers; the outermost initially produced the ground meristem and protoderm, and the innermost produced the procambium. The immature endodermis of the parent root contributed to the early stages of the root tip as an endodermal covering. Prior to emergence, the ground meristem/protoderm produced cells into the endodermal covering. After emergence, the endodermal covering was replaced by a calyptrogen, which was derived from the ground meristem/protoderm and which, in turn, formed the rootcap. A typical monocotyledonous three-tiered meristem was then produced. An outer ground meristem also arose before emergence to form a hypodermis in many lateral roots; in these, crystalliferous cell production began in midcortex cells before emergence, and a small aerenchyma developed in their cortices. The rootcap columella stored small amounts of starch shortly after emergence. Lateral roots of T. glauca were smaller than their parental adventitious roots; they normally had only two to six poles of xylem and phloem, and the cortex was less than six cells across. During 1–3-cm elongation, the lateral root apical meristem and mature regions narrowed, stored starch disappeared, fewer crystals formed, aerenchyma production ceased, and the roots stopped elongation.