Sucrose accumulation in sweet sorghum stem internodes in relation to growth

Sweet sorghum (Sorghum bicolor L. Moench) stems of different cultivars (NK 405. Keller and Tracy) reveal a different pattern of sucrose accumulation with respect to in-ternodal sugar content and distribution. The onset of sucrose storage is not necessarily associated with the reproductive stage of the plant, as was hitherto assumed, but obviously occurs after cessation of internodai elongation as was postulated for the sugarcane stem. For at least two of the three cultivars, ripening is an internode to internode process beginning at the lowermost culm parts. Intensive growth of the internodes, combined with a high hexose content in stern parenchyma, shows a strong positive correlation (r |Mg 0.94) to the activity of sucrose synthase (SuSy; EC 2.4.13), but not to invertase (EC 3.2.1.26) which is not present as soluble (neutral and acid) or cell wall-bound, salt-extractable enzyme in the three culsivars investigated. Sucrose synthase measured in sucrose cleavage and synthesis direction reveals divergent activity rates and sensitivity towards exogenously applied Mg²⁺ ions and pH. SuSy activity is connected to the increase of internodai sucrose content in so far as (1) its decline is a prerequisite for the onset of sucrose accumulation and (2) it remains at a constant low level during sucrose storage. Sucrose phosphate synthase (SPS; EC 2.4.1.14) activity in the sorghum stem is low compared to SuSy and uniformly distributed over all inter-nodes. Only source leaves of sorghum show a considerable SPS activity, but neither stem nor leaf SPS reveal a positive correlation to the increase of internodai sucrose content. Sucrose phosphate phosphatase (SPP; EC 3.1.3.24) amounts lo only 24–30% of the respective SPS activity but follows the same distribution pattern. None of the enzymes under study proves to be responsible for the extent of sucrose storage in the stem, so other phenomena such as transport processes within the stern tissue require further investigation.     

Early Exposure to Ethylene Modifies Shoot Development and Increases Sucrose Accumulation Rate in Sugarcane

Sugarcane varieties (Saccharum spp. hybrids) that accumulate high levels of sucrose at the start of the harvest season are of considerable commercial interest. Our understanding of the factors that contribute to early sucrose accumulation in these varieties is limited. In this study we used the plant hormone ethylene to investigate the relationship between growth and early sucrose accumulation in sugarcane. The sugarcane variety KQ228 was exposed to a low concentration of the ethylene-forming compound 2-chloroethylphosphonic acid (CEPA) for a prolonged duration commencing from shoot emergence. The changes in sucrose accumulation and plant growth were investigated. Results from two glasshouse experiments revealed that the CEPA-treated plants accumulated a significantly higher amount of sucrose in their primary culm 2 and 3½ months post-germination. The treated plants had taller primary culms with many smaller internodes, smaller leaves, and a higher photosynthetic rate. Despite producing smaller internodes, treated culms were comparable in fresh weight and volume to the controls due to the compensating effect of faster internode formation. We identified three factors that may have contributed to the early accumulation of more sucrose in the treated culm: (1) the specific leaf area of young leaves was greater indicating efficient diversion of photoassimilate to sink tissue, (2) internode formation was initiated earlier, and (3) internodes continued to form at a faster rate. Consequently, a greater proportion of the internodes in the treated sugarcane matured earlier and began filling with sucrose sooner. The higher reducing sugar level in the apical region of the culm probably contributed to faster internode development. This coincided with elevated vacuolar and cell wall acid invertase gene expression that increased sucrose turnover in the vacuole and increased apoplastic uptake of reducing sugars. These findings extend our understanding of how some sugarcane varieties can naturally accumulate a high level of sucrose early in the season.     

Sucrose Mobilisation in Sugarcane Stalk Induced by Heterotrophic Axillary Bud Growth

A theoretical high-yield sugarcane biofactory can be idealised as containing culm tissue that functions as a secondary source tissue rather than a sink. To investigate this potential process, heterotrophic axillary bud outgrowth from sugarcane (Saccharum spp. hybrids) setts was used as a model system to demonstrate that sucrose is a mobilisable carbon source. The outgrowth and subsequent biomass accumulation of axillary buds from two-eye setts of mature sugarcane stalks grown in the dark was used to measure carbon mobilisation from sett internode pith tissue. After 42 days growth 99.0 ± 0.72% of sett internode pith sucrose was depleted and 2.66 ± 0.16 g of new tissue accumulated. Comparison with a control treatment in which axillary buds were excised at day zero demonstrated that carbon mobilisation was driven by the accumulation of new biomass. Profiling of soluble carbohydrates (viz. sucrose, glucose and fructose), starch, total soluble protein, total amino nitrogen, free amino acids and total insoluble material showed that the sucrose stored in the sett internode pith was the only available carbon source of sufficient size at day zero for the observed biomass accumulation. Other metabolites mobilised were glucose, fructose and some amino acids, notably isoleucine and leucine that were depleted in shoot treatment setts at day 42.     

STEM ELONGATION OF XANTHIUM PLANTS PRESENTED IN TERMS OF RELATIVE ELEMENTAL RATES

Vegetative Xanthium plants grown under noninductive conditions were marked along the stem with India ink and photographed during three successive days. The relative elemental rates of stem elongation [d(dX/dt)/dX] were estimated for 18 plants between 15 and 18 plastochrons. On the average, only the 8.0 cm terminal part of the stem was elongating in this group of plants. Young internodes were elongating at constant relative elemental rates ([d(dX/dt)/dX] was about 0.2 days^–1); nodal portions of the stem beteween two young internodes were not elongating. Internodes longer than 2 cm displayed an acropetal pattern of elongation in which the basal part of an internode stopped elongating and matured first and the apical portion last. The pattern of elongation of the stem could be best approximated to a set of cascading waterfalls with declining plateaus in the direction of the water flow. The acropetal pattern of individual internode elongation observed in Xanthium was similar to those reported for Helianthus and Phaseolus internode growth.     

How does plant population density affect the biomass of Ravenna grass?

Ravenna grass, Tripidium ravennae (L.) H. Scholz, is known to produce an abundance of biomass, but how plant density affects its biomass potential remains unknown. The objectives were to determine the effects of plant density on biomass yield; plant growth traits; biomass?carbon, nitrogen, and ash concentrations; heating value; nitrogen removal; and sucrose concentration in leaves and culms. The treatments consisted of five plant densities (1,250; 2,500; 5,000; 10,000; and 20,000 plants per hectare) in a randomized complete block design with four blocks. Plots were nonirrigated, unfertilized, and harvested once during the dormant season each year. Data were collected from 2015?2019. Dependent variables that varied with plant population density (p < .05) were biomass yield, number of reproductive culms per plant, reproductive culm diameter, reproductive culm sucrose concentration, and nitrogen removal with biomass. Biomass yield ranged from 5.6 to 16.3 Mg/ha for plant densities of 1,250–20,000 plants per hectare, respectively. Combined over years, nonlinear regression of the data showed the equation for biomass yield to plateau at 16.2 Mg/ha at a plant density of 10,640 plants per hectare. As plant density increased, the number of reproductive culms per plant, culm diameter, and culm sucrose concentration significantly decreased. At 1,250 plants per hectare, the number of reproductive culms per plant, culm diameter, and culm sucrose averaged 70, 10.2 mm, and 63.2 g/kg, respectively. Nitrogen removed with biomass significantly increased as biomass yield increased with plant density. At a density of 10,000 and 20,000 plants per hectare, the amount of nitrogen removed annually in the harvested biomass averaged 88 kg/ha. The data suggest that 10,000 plants per hectare would produce the greatest annual biomass yields; however, research is needed to determine the nutrient requirement for Ravenna grass to sustain biomass production at that density.     

Invertase Activity, Carbohydrate Metabolism and Cell Expansion in the Stem of Phaseolus vulgaris L.

In the stem of Phaseolus vulgaris L. the specific activity ofacid invertase was highest in the most rapidly elongating internode.Activity of the enzyme was very low in internodes which hadcompleted their elongation, in young internodes before the onsetof rapid elongation, and in the apical bud. From shortly afterits emergence from the apical bud the elongation of internode3 was attributable mainly to cell expansion. Total and specificactivities of acid invertase in this internode rose to a maximumat the time of most rapid elongation and then declined. Transferof plants to complete darkness, or treatment of plants withgibberellic acid (GA₃), increased the rate of internode elongationand final internode length by stimulating cell expansion. Bothtreatments rapidly increased the total and specific activitiesof acid invertase in the responding internodes; peak activitiesof the enzyme occurred at the time of most rapid cell expansion. In light-grown plants, including those treated with GA₃, rapidcell and internode elongation and high specific activities ofacid invertase were associated with high concentrations of hexosesugar and low concentrations of sucrose. As cell growth ratesand invertase activities declined, the concentration of hexosefell and that of sucrose rose. In plants transferred to darkness,stimulated cell elongation was accompanied by a rapid decreasein hexose concentration and the disappearance of sucrose, indicatingrapid utilization of hexose. No sucrose was detected in theapical tissues of light-grown plants. The results are discussed in relation to the role of acid invertasein the provision of carbon substrates for cell growth. Key words: Cell expansion, Acid invertase, Hexose, Sucrose, Phaseolus     

Elongation of Stem Internodes in the Japanese Morning Glory (Pharbitis nil) in Relation to Auxin Destruction

The relationship between auxin destruction and stem internode elongation was investigated in the vines of the Japanese morning glory (Pharbitis nil Choisy). In young plants an age-dependent gradient was demonstrated in which the decreasing rate of elongation of older internodes correlated with an increasing ability of such tissue to destroy indoleacetic acid. Fragments of tissue from old internodes when incubated with indoleacetic acid (IAA), destroyed the hormone immediately and rapidly; in contrast, young, rapidly elongating internode tissue destroyed IAA only after a lag of several hours. In older plants the gradient was more erratic towards the middle of the plant but old and young tissue behaved as in young plants, i.e., old internodes destroyed IAA rapidly whereas young internodes did not. It appears reasonable to conclude that cessation of elongation in maturing internodes is brought about by developing an internal environment in which auxin is rapidly destroyed.     

Auxin-induced assimilate translocation in the bean stem (Phaseolus vulgaris L.)

Decapitation of the fully-elongated fourth internode of Phaseolus vulgaris plants resulted in the disappearance from the internode of soluble acid invertase (EC 3.2.1.26). This loss was prevented by local applications to the internode of indol-3yl-acetic acid (IAA) and, at the point of IAA application, the specific activity of the enzyme increased by up to 3 times its initial value within 48 h of treatment. IAA applications stimulated the acropetal translocation to the internode of ¹⁴C-sucrose applied to the subtending (second) trifoliate leaf 30 h after decapitation and the start of the auxin treatment. Labelled assimilates accumulated in the IAA-treated region of the internode. Following decapitation the concentration of hexose sugars in the internode fell and that of sucrose rose substantially, but these trends were reversed by IAA treatment. However, small local accumulations of sucrose occurred at the point of auxin application where tissue concentrations of IAA were greatest (determined using [1-¹⁴C] IAA).Considerable quantities of starch were present in the ground parenchyma of the internodes at the start of the experiment but, in the absence of IAA, this was remobilised within 48 h of decapitation. IAA prevented starch loss at and below its point of application to the internode, but not from more distal tissues. Cambial proliferation, radial growth and lignification were stimulated in and below IAA-treated regions of the internode. These observations are discussed in relation to the hormonal regulation of assimilate translocation in the phloem.     

A quantitative trait locus for reduced culm internode length in barley segregates as a Mendelian gene

Yield losses caused by lodging in cereals can be partially controlled by reducing plant height. A progeny of recombinant inbred lines from a cross of two Japanese barley varieties was used to study the inheritance of culm and culm internode lengths. An unexpected QTL for reduced culm length (qCUL), which affected mainly the length of the third and fourth culm internodes, was contributed by ‘Kanto Nakate Gold’. This QTL was also associated with reduced lodging in two experiments. A near-isogenic line (culm length 62.9–73.4 cm) in an ‘Azumamugi’ background, carrying a chromosome segment containing the qCUL allele from Kanto Nakate Gold, was significantly shorter than its recurrent parent (82.9–89.4 cm). The F₂ generation from the next backcross segregated for plant height in a Mendelian monogenic ratio. The qCUL locus was shown to be tightly linked (1.2 cM) with the codominant STS marker ABG608. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The Cellular Pathway of Short-distance Transfer of Photosynthates and Potassium in the Elongating Stem ofPhaseolus vulgarisL. A Physiological Assessment

Plasmolytic disruption of plasmodesmata interconnecting metaphloemsieve element-companion cell complexes with small and largephloem parenchyma cells in the elongating region of internode2 ofPhaseolus vulgarisL. seedlings did not affect accumulationof phloem-imported¹⁴C-photosynthates and⁸⁶rubidium. The membrane-impermeantdye, 5(6) carboxyfluorescein, loaded into leaf phloem as themembrane-permeant diacetate ester, was found not to move radiallyout of the importing sieve elements in the internode elongationregion. In contrast, the apoplasmic tracer, Calcuofluor White,rapidly moved laterally throughout all tissues of the elongationzone. Hexoses, sucrose and potassium were identified as themain osmotica in internode apoplasmic sap. Label asymmetry in[¹⁴C](fructosyl)sucrose was retained on accumulation by excisedstem segments. Uptake of [¹⁴C]sucrose and⁸⁶rubidium by stemsegments exhibited saturation kinetics. Sucrose uptake was inhibitedby the slowly penetrating sulphydryl reagent, para-chloromercuribenzenesulphonicacid.In vitrorates of sucrose uptake, at apoplasmic concentrations,corresponded to its predictedin vivorate of delivery to thestem ground tissues from mature sieve elements when respiratorylosses were assumed to be confined to the stem phloem. For potassium,the total delivery rate could be accounted for by itsin vitrorateof uptake. Overall, it was concluded that radial transport,in the elongation zone of internode 2 ofPhaseolus vulgarisL.seedlings, follows an apoplasmic route from mature sieve elementsto stem ground tissues.Copyright 1998 Annals of Botany Company PhaseoluLes vulgaris, apoplasm, elongating stem, French bean, photosynthates, potassium, radial transfer, symplasm.     

A new picture of cell wall protein dynamics in elongating cells of <Emphasis Type="Italic">Arabidopsis thaliana</Emphasis>: Confirmed actors and newcomers

Background  

Cell elongation in plants requires addition and re-arrangements of cell wall components. Even if some protein families have been shown to play roles in these events, a global picture of proteins present in cell walls of elongating cells is still missing. A proteomic study was performed on etiolated hypocotyls of Arabidopsis used as model of cells undergoing elongation followed by growth arrest within a short time.     

The AGCVIII kinase Dw2 modulates cell proliferation,endomembrane trafficking,and MLG/xylan cell wall localization in elongating stem internodes of Sorghum bicolor

Stems of bioenergy sorghum (Sorghum bicolor L. Moench.), a drought-tolerant C4 grass, contain up to 50 nodes and internodes of varying length that span 4–5 m and account for approximately 84% of harvested biomass. Stem internode growth impacts plant height and biomass accumulation and is regulated by brassinosteroid signaling, auxin transport, and gibberellin biosynthesis. In addition, an AGCVIII kinase (Dw2) regulates sorghum stem internode growth, but the underlying mechanism and signaling network are unknown. Here we provide evidence that mutation of Dw2 reduces cell proliferation in internode intercalary meristems, inhibits endocytosis, and alters the distribution of heteroxylan and mixed linkage glucan in cell walls. Phosphoproteomic analysis showed that Dw2 signaling influences the phosphorylation of proteins involved in lipid signaling (PLDδ), endomembrane trafficking, hormone, light, and receptor signaling, and photosynthesis. Together, our results show that Dw2 modulates endomembrane function and cell division during sorghum internode growth, providing insight into the regulation of monocot stem development.     

The biophysical basis of elongation growth in internodes of deepwater rice

Partial submergence induces rapid internodal elongation in deepwater rice (Oryza sativa L., cv Habiganj Aman II). We measured in vivo extensibility, tissue tension, hydraulic conductance and osmotic potential in the region of cell elongation in the uppermost internode. The in vivo extensibility of the internode, measured by stretching of living tissue with a custom-made constant stress extensiometer, rose rapidly following submergence of the plant. Both the elastic (E_el) and plastic (E_pl) extensibility increased when growth of the internode was induced. The submerged internode displayed tissue tension (elastic outward bending of longitudinally split internode sections); in air-grown control internodes, no such bending occurred. The hydraulic conductance, estimated from the kinetics of tissue shrinkage in 0.5 molar mannitol and subsequent swelling in distilled water, was not changed by submergence. The osmotic potential, measured with a dew-point hygrometer using frozen-thawed tissue, was only 18% less negative in the submerged internode than in the air-grown control. This indicates that osmoregulation takes place in rapidly elongating rice internodes. We suggest that the rapid expansion of the newly formed internodal cells of submerged plants is controlled by the yielding properties (E_pl) of the cell walls. Experiments with excised stem sections indicate that gibberellin is involved in increasing the E_pl of the elongating cell walls.     

Effects of potassium and gibberellic acid on stem growth of whole sunflower plants

Abstract The effects of gibberellic acid (GA₃) on whole sunflower (Helianthus annuus L.) plants grown at three potassium (K) levels (0.0, 0.5 and 5.0 mM) were studied. A tenfold increase in the length of the first internode was observed when plants grown without K were treated with GA₃. The uneven K distribution along the plant (higher K content in the higher internodes) was enhanced by GA₃ treatment. Gibberellic acid increased the content of reducing sugars, especially in K-deficient plants. An increase in the K level in the nutrient solution resulted in a decrease of the osmotic potential of stem segments. Osmotic potential differences within the elongating first internode were increased by GA₃ treatment.     

The cellular basis of the elongation response in submerged deep-water rice

The cellular basis of internode elongation was studied in intact deep-water rice plants (Oryza sativa L. cv. Habiganj Aman II) and in isolated stem sections. In intact plants, growth was stimulated by submergence in water and by ethylene treatment. In isolated sections, growth was enhanced by submergence, by ethylene, and by exposure of the tissue to an atmosphere of 3% O₂, 91% N₂ and 6% CO₂ or 3% O₂, 91% N₂, 6% CO₂ and 1 l l^-1 C₂H₄ (by vol.). Under all these conditions, growth was localized in the intercalary meristem at the bases of the internodes. Autoradiography of [³H]thymidine-labeled tissue showed activation of cell division and longitudinal expansion of the intercalary meristem. Increased production of new cells and their subsequent elongation thus form the basis for the growth response to submergence and ethylene treatment in deep-water rice plants.     

Characterising the morphological characters and carbohydrate metabolism of oat culms and their association with lodging resistance

• Lodging resistance can be improved by enhancing the mechanical strength of culms, and culm carbohydrates could improve this mechanical strength. Culm carbohydrates can regulate development of the culm and affect its toughness.
• The present study determines the relationship between lodging and carbohydrate content in oat culms. Field experiments were conducted in alpine regions in 2017 and 2018 using three oat varieties with different lodging resistance. Lodging‐related morphological characteristics were directly determined and culm carbohydrate content and enzyme activity related to cellulose synthesis and sucrose metabolism were evaluated with ultraviolet spectrophotometry.
• Results showed that the lower the gravity height or the lower ratio of gravity height to plant height, the stronger the lodging resistance of the varieties. Higher culm nonstructural (NSC) and structural (SC) carbohydrate content contributed to the ability of culms to resist lodging, especially the content of cellulose and sucrose. PCA showed that sucrose metabolism and SC content were closely related to lodging resistance. Correlation analysis showed that the lodging index (LI) was significantly negatively correlated with NSC. Sucrose content was highly and significantly positively correlated with NSC. Additionally, the activities of sucrose phosphate synthase (SPS) and sucrose synthase (SS) were highly and significantly positively correlated with sucrose and cellulose content.
• The relationship between field characters and oat lodging, as well as the regulatory mechanism of carbohydrate content on lodging resistance of the culm are discussed.

     

<Emphasis Type="Italic">BMP-2</Emphasis> gene-fibronectin-apatite composite layer enhances bone formation

Background  

Safe and efficient gene transfer systems are needed for tissue engineering. We have developed an apatite composite layer including the bone morphogenetic protein-2 (BMP-2) gene and fibronectin (FB), and we evaluated its ability to induce bone formation.