REGIONS OF CELL DIVISION AND ELONGATION DURING STEM GROWTH OF XANTHIUM

Regions of cell division and cell elongation were established during stem growth of Xanthium pensylvanicum (cocklebur). From percent mitosis it was determined that the region of cell division in a Xanthium stem of Plastochron Index (PI) 13.89 was 20 mm long, starting from the stem apex and proceeding in a basipetal direction. Measurements of cortical cell lengths demonstrated that mature cell length in the stem of the same plant was reached at about 55 mm distance from the stem apex. Between 20 and 55 mm distance from the apex, shoot growth occurred by cell elongation alone. The maximum rate of change in cell length (dC/dX) was at about 15 mm distance from the stem apex. The length of the apical growth region was a function of the age of the plants. The older the plant, the longer its apical region of elongation. Growth of Xanthium stems was due primarily to elongation of internodes; the nodal regions did not seem to elongate.     

Relation of growth process to spatial patterns of electric potential and enzyme activity in bean roots

The electric spatial pattern and invertase activity distribution in growing roots of azuki bean (Phaseolus chrysanthos) have been studied. The electric potential near the surface along the root showed a banding pattern with a spatial period of about 2 cm. It was found that the enzyme activity has a peak around 3-7 mm from the root tip, in good agreement with the position of the first peak of the electric potential, which is located a little behind the elongation zone. An inhomogeneous distribution of ATP content was also detected along the root. Experiments on the electric isolation of the elongation zone from the mature zone and acidification treatment showed that H+ is transported from the mature-side to elongation-side regions, causing tip elongation through an acid-growth mechanism. Both acidification and electric disturbance on growing roots affected growth significantly. Simultaneous measurements of electric potential and enzyme activity clearly showed a good correlation between these two quantities and growth speed. From an analogy with the Characean banding, the spatio-temporal organization via the cell membrane in electric potential and enzyme activity can be regarded as a dissipative structure arising far from equilibrium. These experimental results can be interpreted with a new mechanism that the dissipative structure is formed spontaneously along the whole root, accompanied by energy metabolism, to make H+ flow into the root tip.     

Growth and electric current loops in plants

A theory is presented for a relationship between ion accumulation and electric current loops in multicellular systems such as the roots and stems of higher plants. A network of electric circuits shows that the electric current transported across the cell membrane flows between an elongating region and a mature region, not only in roots but also in stems. In roots, ions constituting the extracellular electric current flow in the external aqueous medium, while in stems an electric current of comparable density flows within the epidermal cell wall. Based on this theoretical result, electric isolation between the elongating and mature regions was made in the case of both roots and stems. The speed of growth during the initial stage was greatly decreased due to a change in the distribution of protons around the surfaces of the plant by cutting off the electric current loop. Electrochemical calculation shows that ions are not always accumulated at the efflux site, since the ion distribution is strongly affected by the relation of the magnitudes between the electric field and electric current. The results calculated for the electric potential and pH distributions around the root agree with experimental data.     

Band structure of surface electric potential in growing roots

For growing roots of azuki bean (Phaseolus chrysanthos), an electric potential is measured minutely along the surface of the root, together with the surface pH. It was found that the root begins to display a band-type pattern of potential with a spatial period of about 2 cm in a mature region as soon as it grows to about 10 cm in root length, while the surface potential shows only one convex peak around a position 5-20 mm behind a root tip and a succeeding concave peak around 20-35 mm, providing the length of root is shorter than about 10 cm. Since the surface potential takes a relatively positive value on average at the side of the root base compared with that in an elongation zone near the tip, electric current is supposed to flow into the elongation zone, accompanied by some local current loops in the mature region. The present band-type pattern observed first in multi-cell systems seems to be a kind of dissipative structure appearing far from equilibrium, and hence its relationship to growth is discussed.     

Modelling the distribution of diameter growth along the stem in Scots pine

This paper presents an empirical model for the distribution of diameter growth along the stem in Scots pine (Pinus sylvestris L.) and for the consequent stem form over time. First, the distribution of annual mass growth in the stem is determined as a function of the total annual growth in stem mass, current stem mass and the distribution of the latter along the stem. Second, the distribution of diameter growth is obtained by converting the fraction of annual growth in the stem mass at a given height in the stem into the thickness of the annual ring at the same height. Application of the model to Scots pine data sets including both young and mature trees not used in parameter estimation showed that the model was capable of reconstructing the distribution of diameter growth from the stem butt to the apex and from the pith to the stem surface at any height in the stem in both young and mature trees. The resulting empirical model was also linked to a physiological, process-based model in order to study its performance in a simulated stand. Simulations representing trees grown in unthinned and thinned Scots pine stands with trees of different status (from dominant to suppressed) showed that the response in tree growth to thinning in terms of the distribution of diameter growth along the stem was quite realistic relative to measured data.     

Co-regulation Of ear growth and internode elongation in corn

Ear is the harvest part of corn (Zea mays L.) and we are interested in studying its growth and development in our effort in corn yield improvement. In our current study, we examined the relationship between ear growth and internode characteristics using different approaches. Correlations between stem growth rate and number of ears per plant (prolificacy) were assessed among several genotypes. Internode elongation of 2 genotypes was modified by plant hormones and by population density manipulations. Among the 7 genotypes examined that have different prolificacy levels, there was a general correlation of slower stem elongation at middle growth stages and larger ear number. When the internode elongation was enhanced by application of gibberellic acid (GA), ear growth was suppressed; and when a GA synthesis inhibitor uniconazole was applied at early stages, internode length was reduced and ear growth was promoted in terms of both ear size and visible ear number at silking stage. Higher population density caused longer internodes and fewer ears per plant and the effect of lower density was the opposite. Our results suggested that internode elongation in the middle section of corn plants was linked to suppression of ear development in corn.     

Modulation of leaf elongation, tiller appearance and tiller senescence in spring barley by far-red light

Supplemental far-red (FR) illumination of light-grown grass seedlings inhibits tiller production while enhancing leaf elongation. Although much is known about FR enhancement of internode elongation in dicots, relatively little research has been conducted to determine the effects of FR on monocot development. In growth chamber experiments, fibre optics were used to direct supplemental FR to elongating leaf blades, main stem bases and mature leaf blades of light-grown barley (Hordeum vulgare L.) seedlings. Our objective was to identify specific sites of perception for FR enhancement of leaf elongation and inhibition of tiller production, and to assess potential FR effects on tiller senescence. Far-red illumination of elongating leaves or of the main stem base reduced the total number of tillers per plant, primarily by reducing secondary and tertiary tiller production, and enhanced leaf elongation. However, leaf elongation was less sensitive to stem base treatments than to illumination of the elongating blade. Increased leaf length resulted from increased leaf elongation rate, while the duration of leaf elongation was unaffected. Exposure of mature leaf blades to FR had no effect on tillering or leaf elongation. None of the FR treatments led to tiller senescence. Localization of FR perception in vertically oriented tissues such as elongating blades and stem bases permits early detection of reflected light from neighbouring plants, allowing rapid response to impending competition.     

Rapid Changes in the Pattern of Electric Current around the Root Tip of Lepidium sativum L. following Gravistimulation

Using a highly sensitive vibrating electrode, the pattern of naturally occurring electric currents around 1-day-old primary roots of Lepidium sativum L. growing vertically downward and the current pattern following gravistimulation of the root has been examined. A more or less symmetrical pattern of current was found around vertically oriented, downward growing roots. Current entered the root at the root cap, the meristem, and the beginning of the elongation zone and left the root along most of the elongation zone and in the root hair zone. After the root was tilted to a horizontal position, we observed current flowing acropetally at the upper side of the root cap and basipetally at the lower side within about 30 seconds in most cases. After a delay of several minutes, acropetally oriented current was also found flowing along the upper side of the meristematic zone. The apparent density of the acropetal current in the root cap region increased and then decreased with time. Gravitropic curvature was first visible approximately 10 minutes after tilting of the root to the horizontal position. Since the change in the pattern of current in the root cap region precedes bending of the root and is different for the upper and lower side, a close connection is suggested between the current and the transduction of information from the root cap to the elongation zone following graviperception in the cap.     

STEM ANATOMY AND INTERNODAL DEVELOPMENT IN PHASEOLUS VULGARIS

Internodal length in Phaseolus vulgaris decreases slightly from one through three, then sharply increases in four, and subsequently decreases toward the apex. Two internodes undergo rapid growth at a given time, although they do not mature concurrently. Early growth of the internode takes place throughout, with later growth limited to successively higher levels. Elongation results from both cell division and cell elongation, and the relative importance of these two phenomena is significant in determining final length. The difference in final length between internodes is brought about by rate rather than period of growth. The procambium is relatively homogeneous in structure, whereas the cambium consists of both fusiform and ray initials. During stem development the vascular meristem gradually acquires the characteristics of the cambium. By the time elongation ceases, the meristem has developed all the characteristics of the cambium.     

Two distinct regions of response drive differential growth in Vigna root electrotropism

Although exogenous electric fields have been reported to influence the orientation of plant root growth, reports of the ultimate direction of differential growth have been contradictory. Using a high‐resolution image analysis approach, the kinetics of electrotropic curvature in Vigna mungo L. roots were investigated. It was found that curvature occurred in the same root toward both the anode and cathode. However, these two responses occurred in two different regions of the root, the central elongation zone (CEZ) and distal elongation zone (DEZ), respectively. These oppositely directed responses could be reproduced individually by a localized electric field application to the region of response. This indicates that both are true responses to the electric field, rather than one being a secondary response to an induced gravitropic stimulation. The individual responses differed in the type of differential growth giving rise to curvature. In the CEZ, curvature was driven by inhibition of elongation, whereas curvature in the DEZ was primarily due to stimulation of elongation. This stimulation of elongation is consistent with the growth response of the DEZ to other environmental stimuli.     

Relationship between Growth and Electric Oscillations in Bean Roots

Extracellular and intracellular electric potentials in bean roots are known to show electric oscillations along the longitudinal axis with a period of several minutes. The relationship between growth and the electric oscillations was studied using roots of adzuki (Phaseolus chrysanthos). We measured surface electric potentials with a multielectrode apparatus while simultaneously measuring elongation using a CCD camera and monitor. Roots having an electric oscillation grew faster than roots with no oscillation. Furthermore, elongation rate was higher in roots with higher oscillation frequency. Oscillation frequency had a strong dependence on temperature; i.e. Q₁₀ was estimated at 1.7. These results suggest a correlation between electric oscillation and elongation.     

How the scrub height of dwarf pine Pinus pumila decreases at the treeline

Plant height decreases much within narrow altitudinal spans near treelines. We compared the stem age, stem inclination and shoot elongation rates of alpine dwarf pine Pinus pumila between the upper distribution limit (treeline, 2,850 m a.s.l.) and the lower distribution limit (2,500 m a.s.l.) on Mount Norikura in central Japan, to examine how the growth traits of P. pumila change with altitude. The mean stem height at the upper distribution limit (49 cm) was about a quarter of that at the lower distribution limit (187 cm). The mean ratio of stem height to length was lower at the upper distribution limit than at the lower distribution limit, indicating that P. pumila stems inclined more at the higher altitude. The mean stem age at the upper distribution limit (48 years) was less than a half of that at the lower distribution limit (109 years). Although the shoot elongation rate positively correlated with stem length at the two altitudes, the shoot elongation rate at a given stem length was lower at the upper distribution limit than at the lower distribution limit. Thus, less developed scrub at the upper distribution limit than at the lower distribution limit was due to shorter stem age, more creeping stems and lower shoot elongation rates. Generally, wind velocity is greater in higher altitudes. Probably, strong wind reduces the growth and mean stem age of P. pumila stems at the upper distribution limit. Therefore, this study concludes that the scrub height of P. pumila is controlled not only by temperature, but also by strong wind.     

Laser micromarking technique in studying the negative gravitropism in pea stem

A laser micromarking technique on plant epidermis was developed to study how a plant can reduce the stress in bending behavior by controlling the growth and morphogenesis. The negative gravitropism in a pea seedling (Pisum sativum L.) was discussed based on the time-dependent displacement of laser marking points which were formed by spatially-selective laser ablation of the cuticle layer that covers the outer surface of a plant. The elongation of the stem in the horizontal direction was remarkable in the first half of the gravitropism. The elongation percentages of the stem length between laser-marking points at around upper surface, middle, and bottom surface were evaluated to be 2.57, 4.87, and 7.70%, respectively. The characteristic feature of the stem bending in gravitropism is the elongation even at the upper surface region, that is, inside of the bending. This is a different feature from cantilever beams for structural materials like metals and polymers, where the compression of the upper surface and elongation of the bottom surface are caused by bending. Another laser micromarking technique was developed to improve the resolution of a dot-matrix pattern by fluorescent material transfer to a plant through a masking film with a micro-hole matrix pattern. Similar time-dependent displacement behavior was observed for a fluorescent dot-marked stem showing a feedback control loop in the mechanical optimization. These results suggested that plants solve the problem of the stress in stem bending through growth. The laser micromarking is an effective method for studying the mechanical optimization in plants.     

Ion currents and the nitrogen status of roots of Hordeum vulgare and non-nodulated Trifolium repens

Abstract. Profiles of self-generated ion currents associated with the growing primary root tips of intact Hordeum vulgare L. and Trifolium repens L. (nonnodulated) seedlings were measured using a highly sensitive vibrating electrode in media containing NH⁺₄ or NO^-₃, and compared to control roots growing in nitrogen free media. Under these three nutrient regimes, positive current entered the root at regions corresponding to the meristematic tissues and main elongation zones of root tips and left from the mature root tissues. Mapping the surface of the roots with a pH-sensitive microelectrode revealed regions of external alkalinity where positive electrical current entered the root, and external acidity where positive current exited. The correlation between pH-profile and the pattern of ion current generation in these experiments suggests that H⁺ ions were responsible for carrying the bulk of the root-generated current. Assimilation of NHJ results in net H⁺ extrusion while assimilation of NO^-₃, results in net OH^-₃ efflux. Growth on NH⁺₄, as compared to growth on NO^-₃, stimulated the magnitude of the electrical current but did not affect significantly the growth rate of the roots. However, despite the differing stresses on internal pH regulation that arise due to growth on the two exogenous forms of combined nitrogen, the current profiles were qualitatively similar under the different conditions that were examined. The role of the circulating proton current is not yet known; however, the constancy of the current profile under different nutrient regimes sustains the hypothesis that the current may have a role in the regulation of root polarity.     

In vitro regeneration of the tropical multipurpose leguminous tree Sesbania grandiflora from cotyledon explants

Summary A system using cotyledon pieces as explants and a BAP/NAA containing medium was developed for in vitro mass propagation of Sesbania grandiflora, a tropical nitrogen-fixing leguminous tree. The age and the lighting conditions of seedlings providing the explants were shown to be critical factors for both bud induction and bud elongation. Optimal choice for an efficient and reproducible bud induction process consisted of dark-grown seedlings, 24/36 h-old-post-imbibition, that yielded up to 96% of explants producing more than 30 buds each, after one week in culture. Bud development occurred throughout a direct organogenesis pathway, from the proximal and adaxial cut surface of the explants as proved by histological studies. Additional sites of regeneration were also obtained after wounding on the epidermal surface of explants, suggesting a large distribution of regenerative cells all along the explants. Bud elongation, i.e. stem differentiation and leaf growth, was improved by bud isolation from cotyledon explants and their further subculture in liquid bud elongation media for one week. Rooting was obtained on an auxin medium after 3 weeks and plants were established in soil with 92% success.Abbreviations BAP 6-BenzylAminoPurine - NAA a-Naphthalene Acetic Acid - IBA Indole-3-Butyric Acid - IAA Indole-3-Acetic Acid - GA3 Gibberellic Acid - MS Murashige and Skoog (1962) medium     

Persistence of Triazole Growth Retardants on Stem Elongation of Rhododendron and Kalmia

Triazole growth retardant chemicals may inhibit stem elongation of woody ornamental species for several years after application. Potted plants of large-leaf Rhododendron catawbiense and Kalmia latifolia were treated with a single spray application of paclobutrazol or uniconazole in the 2nd year from propagation. They were transplanted into the field the next spring. The elongation of stems was measured in the year of application and in the next 2–4 years. Treatments with a wide range of doses were applied in 1991, 1992, or 1995. For all except the most dilute applications, stem elongation was retarded in the year after application. At the highest doses, stem growth was inhibited for 2 years after application. The results were fit to a model of growth regulator action which assumed that stem elongation was inversely related to the amount of growth regulator applied. For paclobutrazol, the dose per plant that inhibited stem elongation half as much as a saturating dose was tenfold that for uniconazole, about 0.5 and 0.05 mg, respectively. For both chemicals, the dose-response coefficient decreased exponentially with time after application, with an exponential time constant of about 2 year⁻¹. A dose of growth regulator which reduced stem elongation by half immediately after application would only inhibit 12% of stem elongation the next year. However, a tenfold greater dose would result in less than half the stem elongation of untreated plants in the next year. Received February 28, 1997; accepted July 8, 1997     

Stem elongation of ornamental bromeliad in tissue culture depends on the temperature even in the presence of gibberellic acid

Acanthostachys strobilacea Link, Klotzsch, & Otto is an ornamental bromeliad native to Brazilian Atlantic Forest that naturally exhibits a rosette growth pattern. According to the temperature conditions of the in vitro culture, this species can exhibit stem elongation, facilitating the isolation of the nodal segments to be applied in its micropropagation. The rosette morphology is reestablished when this species is maintained under low temperature, thus allowing the maintenance of a germplasm collection (slow growth storage). Gibberellins (GA) are usually applied to stimulate stem elongation in micropropagated plants. Thus, our aim here was to verify the influence of temperature over the stem elongation of A. strobilacea when GA₃ is applied to the medium, thus estimating the use of this phytoregulator in slow growth cultures at low temperatures. Physiological and anatomical studies were performed on plants obtained from nodal segments maintained at 10, 15, 20, and 25 °C. Regardless of the applied treatment, no segments developed at 10 °C. Stem elongation occurred at 25 and 30 °C, and was not seen for plants grown under 15 and 20 °C. The application of 50 µM of GA₃ restored stem elongation in plants at 20 but not at 15 °C. The influence of gibberellins on stem elongation of this tropical bromeliad depends on the cultivation temperature, in which low temperature preponderates over the stem elongation effects of GA₃. In addition, the optimum temperature for the slow growth of this species depends on the starting temperature of the explant used in the micropropagation.     

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.     

Growth enhancement of transgenic poplar plants by overexpression of Arabidopsis thaliana endo-1,4–β-glucanase (cel1)

Poplar (Populus tremula) plants which had been transformed with Arabidopsis thaliana cel1 cDNA and successfully over-expressed the gene, exhibited significant phenotypic alterations which included taller plants, larger leaves, increased stem diameter, wood volume index, dry weight and a higher percentage of cellulose and hemicellulose, compared to the wild-type plants. Transgenic A. thaliana plants over-expressing A. thaliana cel1 exhibited similar levels of cel1 mRNA in the elongation zone of the flowering stem and higher levels in mature leaves when compared with wild-type plants. CEL1 protein levels in the elongation zone of the flowering stem of transgenic plants were similar or slightly higher compared to that of the wild-type plants, whereas mature leaves of transgenic plants contained a higher level of CEL1. These data indicate that in elongating zone of Arabidopsis, CEL1 level is tightly regulated. In contrast to transgenic poplar over-expressing the A. thaliana cel1, no phenotypic difference was found between A. thaliana transgenic and wild-type plants.     

Ethylene and the Regulation of Apple Stem Growth under Stress

Bending stree resulted in an increase in the ethylene concentration in the internal atmosphere of apple stem (Malus domestica Borkh. cv. Winesap). reaching a maximum at about 2 days after bending. The rise in ethylene content was followed by a depression of growth at about 14–21 days. Ethylene content returned to control levels after about 3 weeks. Application of a past naphthaleneacetic acit caused a similar increase in ethylene levels, and the application of ethephon pastes brought about an inhibition of elongation growth. Whereas stress treatment resulted in an inhibition of growth in stem diameter as well as elongation of growth in stem diameter as well as elongation, the ethephon applications resulted in a stimulation of growth in diameter. It is suggested that ethylene may be involved in the growth responses to mechanical stress.