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.     

Developmental Morphology of Roots and root-borne shoots ofPodostemum subulatum as compared withZeylanidium olivaceum (Podostemaceae—Podostemoideae) Part VII of the series ‘Morphology of Podostemaceae’

The root structure ofPodostemum subulatum is investigated and compared with that ofZeylanidium olivaceum. Podostemum has thread- or ribbon-like roots. The root tip consists of an inner apical meristem and a single-layered root cap. From roots arise numerous shoots of endogeneous origin. Their vascular bundle isab initio connected with the root bundle.By the simple (reduced) apical zonation, the roots ofPodostemum subulatum appear more advanced than the crustose roots ofZeylanidium olivaceum, which bear an ordinary (though asymmetrical) root cap. With regard to the endogeneous root-borne shoots, however,Zeylanidium appears more advanced because of the shoot dimorphism. The floriferous shoots have a short axis that grows plagiotropously above the crust surface, whereas the axes of the vegetative shoots are extremely short and remain, together with the apical meristem, within the crust. Only the leaves protrude from the crust surface.     

Comparative developmental anatomy of the root in three species of Cladopus (Podostemaceae)

Root meristem structure and root branching in three species of Cladopus were investigated from developmental and anatomical perspectives. Cladopus fukiensis has a compressed bell-shaped meristem at the apex of a compressed subcylindrical root, while C. javanicus and perhaps C. nymanii, with a ribbon-like root, have a half lozenge-shaped ( subset as seen from above) meristem composed of an apical meristem of cubic cells and a marginal meristem of rectangular cells. The dorsiventrality of the meristem results in root dorsiventrality, and a marginal meristem contributes to the broadening of the root. Comparisons of meristem structure and root morphology suggest that the ribbon-like root of, e.g. C. javanicus, evolved towards the foliose root of Hydrobryum, sister to the genus Cladopus, by loss of an indeterminate apical meristem. The lateral root of C. javanicus initiates within the meristem of a parent root. The dorsal dermal layer and inner cells of the lateral-root meristem appear endogenously under the dermal layer of the parent root, while the ventral layer is derived exogenously from a ventral dermal layer continuous with the parent-root meristem. This mosaic pattern of exogenous and endogenous root formation differs from the truly exogenous formation seen in Hydrobryum and Zeylanidium. The dorsiventral mosaic origin of the root meristem may account for root cap asymmetry.     

Leaf development in the absence of a shoot apical meristem in Zeylanidium subulatum (Podostemaceae)

* BACKGROUND AND AIMS: The Podostemaceae are a family of unusual aquatic angiosperms that live in rapids and waterfalls. To adapt to such extreme habitats, the family shows unusual morphologies. This study investigated the developmental anatomy of the shoot of Zeylanidium subulatum borne on the prostrate root attached to submerged rock surfaces. * METHODS: Shoots of Z. subulatum were observed under the microscope using resin-sections. * KEY RESULTS: The shoot has no shoot apical meristem (SAM) and, without it, forms leaves distichously dorsiventrally facing the immediately older leaf. A new leaf forms on the adaxial side of a pre-existing leaf and also on the abaxial side of a leaf on flowering shoots. In both cases, the young leaf is endogenous below the older leaf and maintains histological continuity with it. Shortly after internal initiation, the leaf primordia become separate from each other due to cleavage between adjacent leaves of opposite ranks. The cleavage is caused by intercellular separation as well as by degeneration of vacuolated cells. Loss of the SAM is probably linked with the speculated shift of the site of leaf formation to the root. * CONCLUSIONS: The 'shoot' of Z. subulatum is characterized by the absence of a SAM, endogenous leaf formation in the absence of a SAM, cleavage between leaf primordia, and adventitious leaf formations. These innovations occur in some Podostemaceae that have become increasingly adapted to extreme aquatic habitats.     

Developmental morphology of foliose shoots and seedlings of Dalzellia zeylanica (Podostemaceae) with special reference to their meristems

The developmental morphology of seedlings and shoots of Dalzellia zeylanica was examined with reference to the meristem in order to understand the dorsiventral, foliose shoot. In seedlings, no obvious primary shoot and no root are formed. Subsequent to disappearance of the vestigial primary shoot meristem, two shoot meristems are established in the axils of the cotyledons, one of which grows into a secondary shoot. Microtome and SEM examinations of mature plants show that the shoot meristem is complex, comprising three zones along the shoot margin. The organogenetic zone, equivalent to the shoot apical meristem, produces dorsal leaves proximally and much fewer marginal leaves distally. During development, the zone repeatedly changes into a dorsal zone, while a new organogenetic zone is formed in an area between developing marginal leaves, resulting in the alternation of the organogenetic and dorsal zones, which allowed development of the coenosomic structure of the shoot. The dorsal and ventral zones do not produce leaves, but contribute to shoot expansion. The ventral zone also forces the marginal leaves to shift to the lateral side of the shoot. The rosette with tufted leaves might be a modification of the short shoot (ramulus) of other Tristichoideae.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 144 , 289–302.     

Comparative anatomy of embryogenesis in three species of Podostemaceae and evolution of the loss of embryonic shoot and root meristems

During embryogenesis in angiosperms, the embryonic shoot and root meristems are created at opposite poles of the embryo, establishing a vertical body plan. However, the aquatic eudicot family Podostemaceae exhibits an unusual horizontal body plan, which is attributed to the loss of embryonic shoot and root meristems. To infer the embryogenetic changes responsible for the loss of these meristems, we examined the embryogenesis of three podostemads with different meristem characters, that is, Terniopsis brevis with distinct shoot and root meristems, Zeylanidium lichenoides with reduced shoot and no root meristems, and Hydrobryum japonicum with no shoot and no root meristems. In T. brevis, as in other eudicots, the putative organizing center (OC) and L1 layer (=the epidermal cell layer) arose to generate a distinct shoot meristem initial, and the hypophysis formed the putative quiescent center (QC) of a root meristem. Z. lichenoides had a morphologically unrecognizable shoot meristem, because a distinct L1 layer did not develop, whereas the putative OC precursor arose normally. In H. japonicum, the vertical divisions of the apical cells of eight-cell embryo prevented putative OC initiation. In Z. lichenoides and H. japonicum, the putative QC failed to initiate because the hypophysis repeated longitudinal divisions during early embryogenesis. Based on their phylogenetic relationships, we infer that the conventional embryonic shoot meristem was lost in Podostemaceae via two steps, that is, the loss of a distinct L1 layer and the loss of the OC, whereas the loss of the embryonic root meristem occurred once by misspecification of the hypophysis.     

Variation among and within clones in formation of roots and shoots during micropropagation of Campanula isophylla

It is known that treatments enhancing shoot formation often suppress root formation and vice versa. It would be of interest to know if such negative correlations between formation of roots and shoots were also present among genetically different plants, given the same treatment, to ensure that selection for superior shoot formation would not lead to inadvertent decreases in the capacity for root formation. Height and dry weight of micropropagated shoot clusters and the numbers of shoots and roots were measured in 95 seedling clones. Within clones, shoot size was negatively correlated with number of shoots and positively correlated with number of roots. Among clones, however, the number of shoots was not correlated with the size of shoots, but positively correlated with the number of roots. While it is difficult to devise treatments that simultaneously optimize the initiation of roots and shoots, it is thus possible to select for fast-growing clones without compromising root formation.Abbreviations CM clonal means - DCM deviation from clonal means     

Effects of suboptimal root zone temperatures and shoot demand on net translocation of micronutrients from the roots to the shoot of maize

The effects of suboptimal root zone temperatures (RZTs) on net translocation rates from the roots to the shoots and the concentrations of Fe, Mn, Zn, and Cu were examined in maize grown in nutrient solution or soil. Plants were grown at 12 °C, 18 °C and 24 °C RZT. At each RZT, the growth-related shoot demand for nutrients was varied by independently modifying the temperature of the shoot base (SBT) including the apical shoot meristem. The net translocation rates of Mn and Zn from the roots to the shoots were reduced at low RZTs, irrespective of the SBT and of the substrate (soil or nutrient solution). Obviously, the net translocation rates of Mn and Zn at low RZT were mainly regulated by temperature effects on the roots and not by the chemical nutrient availability in the rhizosphere or by shoot growth rate as controlled by SBTs. When both RZT and SBT were reduced, the decrease in net translocation rates of Mn and Zn was similar to the decline in the shoot growth rate and concentrations of Mn and Zn in the shoot fresh matter were not greatly affected or were even increased by low RZT. However, at high SBT and low RZT in nutrient solution, the depressed net translocation rates of Mn and Zn combined with the increased shoot growth resulted in significantly decreased concentrations of Mn and Zn in the shoot, indicating that Mn and Zn may become deficient even at high chemical availability. By contrast to Mn and Zn, the net translocation rates of Fe and Cu at all RZTs were markedly enhanced by increased SBTs. Accordingly, the concentrations of Fe and Cu in the shoot fresh matter were not greatly affected by RZTs, irrespective of the SBTs. These results indicate that the ability of roots to supply Fe and Cu to the shoot was internally regulated by the growth related shoot demand per unit of roots. Deceased 21 September 1996 Deceased 21 September 1996     

Above-ground competition does not alter biomass allocated to roots in Abutilon theophrasti

We tested whether plants allocate proportionately less biomass to roots in response to above-ground competition as predicted by optimal partitioning theory. Two population densities of Abutilon theophrasti were achieved by planting one individual per pot and varying spacing among pots so that plants in the two densities experienced the same soil volume but different degrees of canopy overlap. Density did not affect root:shoot ratio, the partitioning of biomass between fine roots and storage roots, fine root length, or root specific length. Plants growing in high density exhibited typical above-ground responses to neighbours, having higher ratios of stem to leaf biomass and greater leaf specific area than those growing in low density. Total root biomass and shoot biomass were highly correlated. However, storage root biomass was more strongly correlated with shoot biomass than was fine-root biomass. Fine-root length was correlated with above-ground biomass only for the small subcanopy plants in crowded populations. Because leaf surface area increased with biomass, the ratio between absorptive root surface area and transpirational leaf surface area declined with plant size, a relationship that could make larger plants more susceptible to drought. We conclude that A. theophrasti does not reallocate biomass from roots to shoots in response to above-ground competition even though much root biomass is apparently involved in storage and not in resource acquisition.     

Mediation of high-temperature injury by roots and shoots during reproductive growth of wheat

Abstract Previous studies suggest that high temperature stress on wheat (Triticum aestivum L.) involves root processes and acceleration of monocarpic senescence. Physiological changes in wheat roots and shoots were investigated to elucidate their relationship to injury from elevated temperatures after anthesis. Plants were grown under uniform conditions until 10 d after anthesis, when shoot/root regimes of 25°C/25°C, 25°C/35°C, 35°C/25°C and 35°C/35°C were imposed. Growth and senescence of shoots and grain were influenced more by root temperatures than by shoot temperatures. High root temperatures increased activities of protease and RNasc enzymes, and loss of chlorophyll, protein and RNA from shoots, whereas low root temperatures had opposite effects. High root temperatures appeared to induce shoot senescence directly. High shoot temperatures probably disrupted root processes, including export of cytokinins, and induced high leaf protease activity, senescence and cessation of grain development. The authors concluded that responses of wheat to high temperatures, whether of roots or shoots, are manifested as acceleration of senescence and may be mediated by roots during grain development.     

The formation,morphology and anatomy of cluster root of Lupinus albus L. as dependent on soil type and phosphorus supply

The effect of phosphorus supply on the formation, morphology and anatomy of cluster roots of Lupinus albus L. cv Ultra grown in a loam and two sandy soils was examined relative to its effect on total root length, shoot weight and the phosphorus concentration of the shoots. The loam soil was most conducive to the formation of cluster roots. Cluster roots growing in the sandy soils developed to a lesser extent on plants of an equivalent phosphorus status, suggesting that some biotic or abiotic factors independent of phosphorus supply were also operating. The presence of mature cluster rootlets on a length of lateral root increased the root surface area by 14–22 times of an equal length of lateral roots not bearing cluster rootlets. The application of phosphorus decreased cluster-root length, whereas total root length showed a steady increase. There was an inverse relationship between cluster-root production and phosphorus concentration in shoots ranging from 2 to 8.5 mg g^–1 with the critical phosphorus level for maximum shoot growth being around 2.5 mg g^–1. Cluster roots formed in solution culture were not well developed in comparison with those grown in the loam soil or nutrient solution with added loam soil. The organisation of the cluster rootlet was similar to that of the lateral roots. Mature rootlets lacked an apical meristem and a vascular cambium with a reduced root cap and cortical tissue.     

Developmental anatomy of the reproductive shoot in <Emphasis Type="Italic">Hydrobryum japonicum</Emphasis> (Podostemaceae)

Podostemaceae are unusual aquatic angiosperms adapting to extreme habitats, i.e., rapids and waterfalls, and have unique morphologies. We investigated the developmental anatomy of reproductive shoots scattered on crustose roots of Hydrobryum japonicum by scanning electron microscopy and using semi-thin serial sections. Two developmental patterns were observed: bracts arise either continuously from an area of meristematic cells that has produced leaves, or within differentiated root ground tissue beneath, and internal to, leaf base scars after an interruption. In both patterns, the bract primordia arise endogenously at the base of youngest bracts in the absence of shoot apical meristem, involving vacuolated-cell detachment to each bract separately. The different transition patterns of reproductive shoot development may be caused by different stages of parental vegetative shoots. The floral meristem arises between the two youngest bracts, and is similarly accompanied by cell degeneration. In contrast, the floral organs, including the spathella, arise exogenously from the meristem. Bract development, like vegetative leaf development, is unique to this podostemad, while floral-organ development is conserved.     

Carbon allocation to shoots and roots in relation to nitrogen supply is mediated by cytokinins and sucrose: Opinion

In this paper we firstly show some general responses of biomass partitioning upon nitrogen deprivation. Secondly, these responses are explained in terms of allocation of carbon and nitrogen, photosynthesis and respiration, using a simulation model. Thirdly, we present a hypothesis for the regulation of biomass partitioning to shoots and roots.Shortly after nitrogen deprivation, the relative growth rate (RGR) of the roots generally increases and thereafter decreases, whereas that of the shoot decreases immediately. The increased RGR of the root and decreased RGR of the shoot shortly after a reduction in the nitrogen supply, cause the root weight ratio (root weight per unit plant weight) to increase rapidly.We showed previously that allocation of carbon and nitrogen to shoots and roots can satisfactorily be described as a function of the internal organic plant nitrogen concentration. Using these functions in a simulation model, we analyzed why the relative growth rate of the roots increases shortly after a reduction in nitrogen supply. The model predicts that upon nitrogen deprivation, the plant nitrogen concentration and the rate of photosynthesis per unit plant weight rapidly decrease, and the allocation of recently assimilated carbon and nitrogen to roots rapidly increases. Simulations show that the increased relative growth rate of the root upon nitrogen deprivation is explained by decreased use of carbon for root respiration, due to decreased carbon costs for nitrogen uptake. The stimulation of the relative growth rate of the root is further amplified by the increased allocation of carbon and nitrogen to roots. Using the simple relation between the plant nitrogen concentration and allocation, the model describes plant responses quite realistically.Based on information in the literature and on our own experiments we hypothesize that allocation of carbon is mediated by sucrose and cytokinins. We propose that nitrogen deprivation leads to a reduced cytokinin production, a decreased rate of cytokinin export from the roots to the shoot, and decreased cytokinin concentrations. A reduced cytokinin concentration in the shoot represses cell division in leaves, whereas a low cytokinin concentration in roots neutralizes the inhibitory effect of cytokinins on cell division. A reduced rate of cell division in the leaves leads to a reduced unloading of sucrose from the phloem into the expanding cells. Consequently, the sucrose concentration in the phloem nearby the expanding cells increases, leading to an increase in turgor pressure in the phloem nearby the leaf's division zone. In the roots, cell division continues and no accumulation of sugars occurs in dividing cells, leading to only marginal changes in osmotic potential and turgor pressure in the phloem nearby the root's cell division zone. These changes in turgor pressure in the phloem of roots and sink leaves affect the turgor pressure gradients between source leaf-sink leaf and source leaf-root in such a way that relatively more carbohydrates are exported to the roots. As a consequence RWR increases after nitrogen deprivation. This hypothesis also explains the strong relationship between allocation and the plant nitrogen status.     

Enhanced formation of aerenchyma and induction of a barrier to radial oxygen loss in adventitious roots of Zea nicaraguensis contribute to its waterlogging tolerance as compared with maize (Zea mays ssp. mays)

Enhancement of oxygen transport from shoot to root tip by the formation of aerenchyma and also a barrier to radial oxygen loss (ROL) in roots is common in waterlogging‐tolerant plants. Zea nicaraguensis (teosinte), a wild relative of maize (Zea mays ssp. mays), grows in waterlogged soils. We investigated the formation of aerenchyma and ROL barrier induction in roots of Z. nicaraguensis, in comparison with roots of maize (inbred line Mi29), in a pot soil system and in hydroponics. Furthermore, depositions of suberin in the exodermis/hypodermis and lignin in the epidermis of adventitious roots of Z. nicaraguensis and maize grown in aerated or stagnant deoxygenated nutrient solution were studied. Growth of maize was more adversely affected by low oxygen in the root zone (waterlogged soil or stagnant deoxygenated nutrient solution) compared with Z. nicaraguensis. In stagnant deoxygenated solution, Z. nicaraguensis was superior to maize in transporting oxygen from shoot base to root tip due to formation of larger aerenchyma and a stronger barrier to ROL in adventitious roots. The relationships between the ROL barrier formation and suberin and lignin depositions in roots are discussed. The ROL barrier, in addition to aerenchyma, would contribute to the waterlogging tolerance of Z. nicaraguensis.     

Abscisic acid accumulation in the roots of nutrient-limited plants: its impact on the differential growth of roots and shoots

We describe the involvement of abscisic acid (ABA) in the control of differential growth of roots and shoots of nutrient limited durum wheat plants. A ten-fold dilution of the optimal concentration of nutrient solution inhibited shoot growth, while root growth remained unchanged, resulting in a decreased shoot/root ratio. Addition of fluridone (inhibitor of ABA synthesis) prevented growth allocation in favour of the roots. This suggests the involvement of ABA in the redirecting of growth in favour of roots under limited nutrient supply. The ABA content was greater in shoots and growing apical root parts of starved plants than in nutrient sufficient plants. Accumulation of ABA in shoots of nutrient deficient plants was linked to a decrease in leaf turgor. Increased flow of ABA in the phloem apparently contributed to the accumulation of ABA in the apical part of the roots. Thus, partitioning of growth between roots and shoots of wheat plants limited in mineral nutrients appears to be modulated by accumulation of ABA in roots. This ABA may originate in the shoots, where its synthesis is stimulated by the loss of leaf turgor.     

树木根系碳分配格局及其影响因子

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

Developmental anatomy of the shoot apical cell,rhizophore and root ofSelaginella uncinata

The developmental anatomy of the shoot apex, rhizophore and root ofSelaginella uncinata was examined by the semi-thin section method. The shoot apex has a single, lens-shaped apical cell with two cutting faces. Rhizophore primordia are initiated exogenously at the branching point of the second youngest lateral shoot. The rhizophore apex has a tetrahedral apical cell with three cutting faces. A pair of root primordia is initiated endogenously from inner cells of the rhizophore apex, after the rhizophore apical cell becomes unidentifiable losing its activity, and subsequently a root cap is formed from the distal face of the root apical cell. During the course of successive root branching the apical cell in an original root apical meristem becomes unidentifiable and then a new apical cell is initiated in each of the bifurcated root apical meristems. The root branching mode seems to be equivalent to the described dichotomous branching mode of fern shoots. Our results demonstrate a distinct morphogenetical difference between the rhizophore and the root, and confirm the exogenous origin of the rhizophore, as described for other species ofSelaginella. This evidence indicates that the rhizophore is not an aerial root but a leafless, root-producing axial organ.     

Shoot phytochrome B modulates reactive oxygen species homeostasis in roots via abscisic acid signaling in Arabidopsis

Underground roots normally reside in darkness. However, they are often exposed to ambient light that penetrates through cracks in the soil layers which can occur due to wind, heavy rain or temperature extremes. In response to light exposure, roots produce reactive oxygen species (ROS) which promote root growth. It is known that ROS‐induced growth promotion facilitates rapid escape of the roots from non‐natural light. Meanwhile, long‐term exposure of the roots to light elicits a ROS burst, which causes oxidative damage to cellular components, necessitating that cellular levels of ROS should be tightly regulated in the roots. Here we demonstrate that the red/far‐red light photoreceptor phytochrome B (phyB) stimulates the biosynthesis of abscisic acid (ABA) in the shoots, and notably the shoot‐derived ABA signals induce a peroxidase‐mediated ROS detoxification reaction in the roots. Accordingly, while ROS accumulate in the roots of the phyb mutant that exhibits reduced primary root growth in the light, such an accumulation of ROS did not occur in the dark‐grown phyb roots that exhibited normal growth. These observations indicate that mobile shoot‐to‐root ABA signaling links shoot phyB‐mediated light perception with root ROS homeostasis to help roots adapt to unfavorable light exposure. We propose that ABA‐mediated shoot‐to‐root phyB signaling contributes to the synchronization of shoot and root growth for optimal propagation and performance in plants.     

Effect of benzylaminopurine on in vitro and in vivo root development in lentil, Lens culinaris Medik

The effect of benzylaminopurine (BAP) on the formation of roots from lentil shoots regenerated on media containing BAP was studied. Seedling shoot tips, first nodes and bractlets, and immature seeds cultured on the initiation media containing 2.25 or 0.225 mg/l of BAP regenerated multiple bud shoots. The regenerated shoots formed roots in percentages ranging from 4.6 to 39.9% on a rooting medium (R medium) containing 2 mg/l of indoleacetic acid. Rooting success on R medium depended upon the cytokinin used in the initiation media, its concentration, and the time elapsed during shoot formation on these media prior to transplanting regenerated shoots to R medium. In vivo study of root growth of lentil seedlings demonstrated the strong inhibitory effect of BAP on root growth reflected in a drastic reduction of the mitotic index of the root meristem. Received: 27 August 1996 / Revision received: 12 December 1996 / Accepted: 15 January 1997