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.     

Distribution of Three Auxin Protector Substances in Seeds and Shoots of the Japanese Morning Glory (Pharbitis nil)

The existence of substances which inhibit the enzymatic destruction of auxin in shoots of the Japanese morning glory (Pharbitis nil Choisy) has been confirmed, as has the fact that these substances are distributed in a gradient diminishing from apex to base in a manner indicating a regulatory role in internode elongation and tissue maturation. In addition to the 2 auxin protector substances reported previously (protectors I and II) which appear to account for most of the inhibition of the enzymatic destruction of auxin in young, elongating stem tissue, a third substance, designated as protector A, has been found to be highly active in seeds, and shoot tips of mature plants: In germinating seeds, no protector I or II activity was observed; in stem tips, no protector II and only slight protector I activity was observed. In contrast, old tissue contained no detectable amounts of protector A, but did contain protectors I and II. Between these extremes along the shoot axis, mixtures of the 3 substances were found. The evidence can be interpreted to mean that protector A is degraded into protectors I and II and perhaps translocated in this form. Gel filtration studies indicate that protector A has a molecular weight exceeding 200,000 gm/mole.     

Effect of Gibberellic Acid and Cycocel on Tryptophan Metabolism and Auxin Destruction in the Sunflower Seedling

A comparative study of tryptophan conversion in different regions of the sunflower seedling indicates that the regions most active in converting tryptophan on a pathway to auxin are the root apical segments and young leaves; next highest in activity is the cotyledonary tissue. The stem apex proper with leaf primordia is less active than the above regions in converting the auxin precursor. Hypocotyl tissue was observed to be least active. Pre-treatment of the apical bud region of the stem with gibberellic acid (GA) gives rise to tryptophan conversion rates which are 2.1 times those in untreated seedlings. The enhanced tryptophan conversion in the apical bud is followed by an increased elongation rate of the 1st internode which is 2.2 times that in the 1st internode of untreated seedlings. Treatment of the seedlings with Cycocel [(2-chloroethyl)trimethylamnionium chloride] does not reduce tryptophan conversion in the apical bud region of the seedling although elongation of the stem is greatly retarded. Indoleacetic acid (IAA) destruction in cell free preparations as well as in whole sections of the elongating region of the seedling stem was studied. IAA-1-¹⁴C destruction rates with the release of ¹⁴CO₂ in whole sections of 1st internode tissue were approximately 3 times those in cell free preparations of the same region. No significant changes in IAA destruction rates in seedlings pre-treated with GA or Cycocel were observed.     

DNA rearrangements at the region of the dihydroflavonol 4-reductase gene for flower pigmentation and incomplete dominance in morning glory carrying the mutable flaked mutation

 The a-3 ^flecked [J] variegated line of Japanese morning glory bearing white flowers with normal-colored flecks and sectors has been shown to carry a 6.4-kb transposable element, Tpn1, inserted within the DFR-B gene, one of the anthocyanin biosynthesis genes encoding dihydroflavonol 4-reductase (DFR). The a ^flaked [M] variegated line of morning glory also bears white flowers with normal-colored flakes and sectors, and it was shown to carry multiple DNA rearrangements, including insertions of mobile element-like sequences, MELSIP1 and MELSIP2, in its DFR gene region. Unlike the a-3 ^flecked [J] mutation, the mutable a ^flaked [M] allele exhibited incomplete dominance. Interestingly, not only intensely colored flakes but also white spots and sectors were often observed in lightly colored flowers of morning glory in the heterozygous state A[M]/a ^flaked [M]. The interspecific F₁ hybrids between Japanese morning glory and morning glory carrying both a-3 ^flecked [J]/A-3[M] and A[J]/ a ^flaked [M] in the heterozygous condition bear lightly colored flowers with intensely colored sectors as well as white flakes. The results clearly demonstrated that the DFR gene in the a ^flaked [M] line of morning glory is active and complements the DFR-B gene carrying Tpn1 in the a-3 ^flecked [J] line of Japanese morning glory. Interspecific allelic interactions between the mutable a ^flaked [M] gene of morning glory and the corresponding wild-type A[J] gene of Japanese morning glory resulted in incomplete dominance and the formation of white flakes and sectors. The appearance of the white flakes may be due to a somatic mutation of the A[J] gene. Received: 4 November 1996/Accepted: 13 December 1996     

The role of a class III gibberellin 2‐oxidase in tomato internode elongation

A network of environmental inputs and internal signaling controls plant growth, development and organ elongation. In particular, the growth‐promoting hormone gibberellin (GA) has been shown to play a significant role in organ elongation. The use of tomato as a model organism to study elongation presents an opportunity to study the genetic control of internode‐specific elongation in a eudicot species with a sympodial growth habit and substantial internodes that can and do respond to external stimuli. To investigate internode elongation, a mutant with an elongated hypocotyl and internodes but wild‐type petioles was identified through a forward genetic screen. In addition to stem‐specific elongation, this mutant, named tomato internode elongated ‐1 (tie‐1) is more sensitive to the GA biosynthetic inhibitor paclobutrazol and has altered levels of intermediate and bioactive GAs compared with wild‐type plants. The mutation responsible for the internode elongation phenotype was mapped to GA2oxidase 7, a class III GA 2‐oxidase in the GA biosynthetic pathway, through a bulked segregant analysis and bioinformatic pipeline, and confirmed by transgenic complementation. Furthermore, bacterially expressed recombinant TIE protein was shown to have bona fide GA 2‐oxidase activity. These results define a critical role for this gene in internode elongation and are significant because they further the understanding of the role of GA biosynthetic genes in organ‐specific elongation.     

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.     

The possible roles of nutrient deprivation and auxin repression in apical control

Apical control is the suppression of growth in lower branches by a higher dominant branch or leader shoot. We investigated possible mechanisms involved in this developmental response in three widely diverse species (Japanese morning glory, Ipomoea nil, hybrid poplar, Populus trichocarpa, × P. deltoides, and Douglas-fir, Pseudotsuga menziesii). The following two hypotheses were tested: (1) the mineral nutrient-deprivation hypothesis, which is that the continued growth of the lower branches is repressed by the diversion of nutrients to the upper dominating branch or shoot, and (2) the auxin-repression hypothesis, which is that auxin produced in the upper dominating branch or shoot moves down to the lower branches where continued growth is repressed. The results of experiments involving the manipulation of available nutrients by dominant branch removal and fertilization were consistent with the first hypothesis for morning glory, poplar, and for second- or third flushing of lateral branches in Douglas-fir. The results of the experiments involving auxin (NAA, 1-naphthalene acetic acid) replacement treatments on decapitated shoots bearing growing lateral branches were inconsistent with the second hypothesis in morning glory, poplar and in first-flushing Douglas-fir. However, despite concerns about possible NAA toxic effects, there was evidence of auxin repression of second flushing in Douglas-fir. Overall, the data supported a significant role for nutrient availability but not for auxin repression in apical control of morning glory and poplar. In Douglas-fir, apical control in first-flushing lateral branches from over-wintered buds was largely insensitive to both nutrient availability and auxin repression; however, second flushing was sensitive to both.     

水稻茎伸长生长与植物激素

赤霉素（GA）、生长素（IAA）、脱落酸（ABA）和乙烯影响水稻茎（或节间）的伸长,其中赤霉素与水稻茎伸长生长的关系最密切。GA1是植物体内刺激茎伸长的至关重要的赤霉素, GA3已作为最常用的外源激素诱导水稻的节间伸长。水稻茎秆的伸长受激素浓度和敏感性的双重控制,激素浓度或敏感性任一方的改变都有可能导致株高的变异。赤霉素如此显著地促进茎的伸长可能与增加细胞分裂和促使细胞壁松弛有关。而生长素主要促进细胞伸长。植物激素促进水稻茎伸长的分子机理的研究已有较大的进展,预期这方面的研究和应用在未来几年内将有新的突破。     

Action du CCC et du B 995 sur la mise à fleur d'une plante bisannuelle,l'Œnothera biennis

Summary Oenothera biennis L. is a typical biennial plant with an absolute cold requirement for subsequent floral initiation under long-day conditions. Flowering of vernalized Oe. biennis is associated with transition from a rosette habit to the formation of a long flower-bearing stem. Vernalization in Oe. biennis consists of two consecutive stages: (1) preparation for flowering; (2) preparation of stem elongation. At the end of the second stage, the level of endogenous gibberellin-like substances is sufficient to allow the stem elongation which is requisite for subsequent floral development.Applications of (2-chloroethyl)trimethylammonium chloride (CCC) via the roots, when made after the cold treatment, have no effect on internodal elongation and subsequent flowering of vernalized Oe. biennis. Relatively small amounts of CCC applied onto the apical bud of the rosette promote stem growth. However, 750 g of CCC act as a growth inhibitor and delay the beginning of stem elongation but this retardant has no effect on the number of flower buds.In contrast, N-dimethylaminosuccinamic acid (B 995), when applied after a cold treatment, delays the beginning of stem growth, this delay increasing with greater amounts of applied B 995. However, with relatively small amounts of B 995, the rate of later stem growth increases rapidly so that treated plants flower at the same time as controls.If gibberellic acid (GA₃) is applied on the apical bud just at the same time as B 995, it reverses the effects of the growth retardant, even when the amount of B 995, applied by itself, is sufficient to inhibit entirely stem elongation and flowering.B 995 is particularly effective if the preceding cold treatment was just sufficient for effective vernalization. If the cold treatment was extended beyond this time, a greater amount of retardant was required to obtain the same degree of stem growth inhibition.B 995 probably acts by interference with the metabolism of endogenous gibberellin-like substances. The delay of floral initiation in vernalized Oe. biennis by B 995 is a consequence of the retarded internode elongation; if this retardation exceeds a certain limit, the plant is devernalized and exhibits a cold requirement identical with the original one.     

Thermomorphogenic and Photomorphogenic Control of Stem Elongation in Fuchsia Is Not Mediated by Changes in Responsiveness to Gibberellins

Stem elongation in Fuchsia × hybrida was influenced by cultivation at different day and night temperatures or in different light qualities. Internode elongation of plants grown at a day (25°C) to night (15°C) temperature difference (DIF+10) in white light was almost twofold that of plants grown at the opposite temperature regime (DIF−10). Orange light resulted in a threefold stimulation of internode elongation compared with white light DIF−10. Surprisingly, internode elongation in orange light was similar for plants grown at DIF−10 and DIF+10. Flower development was accelerated at DIF−10 compared with DIF+10 in both white and orange light. To examine whether the effects of DIF and light quality on shoot elongation were related to changes in gibberellin metabolism or plant sensitivity to gibberellins (GAs), the stem elongation responses of paclobutrazol-treated plants to applied gibberellins were determined. In the absence of applied gibberellins paclobutrazol (>0.32 μmol plant⁻¹) strongly retarded shoot elongation. This inhibition was nullified by the application of about 10–32 nmol of GA₁, GA₄, GA₉, GA₁₅, GA₁₉, GA₂₀, GA₂₄, or GA₄₄. The results are discussed in relation to possible effects of DIF and light quality on endogenous gibberellin levels and gibberellin sensitivity of fuchsia and their effects on stem elongation. Received October 4, 1997; accepted December 17, 1997     

Day/night temperature environment affects cell elongation but not division in Lilium longiflorum Thunb

Lilium tongiflorum Thunb. cv. ‘Nellie White’ plantswere grown in different day/night temperature (DT/NT) environmentsto determine the anatomical basis for differential responsesof stem elongation to DT and NT. Lilium plants were forced in1986 and 1987 under 25 and 12 different DT/NT environments,respectively, with temperatures ranging from 14 to 30 °C.Parenchyma and epidermal cell length and width were measuredin stem tissue (1987) and epidermal cell length and width weremeasured in leaf tissue (1986). Total cell number per internodeand vertical cell number per internode were calculated. Stemparenchyma and stem and leaf epidermal cell length increasedlinearly as the difference (DIF) between DT and NT increased(DIF = DT —- NT), i.e. as DT increased relative to NT.DIF had no effect on stem parenchyma width, stem and leaf epidermalcell width, or cell number per internode. Data suggested thatstem elongation responses to DIF are elicited primarily througheffects on cell elongation and not division. Key words: Thermoperiodism, thermomorphogenesis, stem elongation, DIF, cell division, cell elongation, leaf expansion     

Promotion of elongation and acid invertase activity in Phaseolus vulgaris L. internode segments by phenylacetic acid

Phenylacetic acid (PAA) significantly stimulated the elongation of isolated Phaseolus vulgaris internodal segments and prevented the decline in acid invertase specific activity observed in segments incubated in the absence of growth substances. Unlike IAA, which stimulated both elongation and invertase activity over a very wide range of concentrations (<10^-4 - 1 mol.m^-3; optimum 10^-2 mol.m^-3), the response to PAA was restricted to a much narrower range of concentrations (3 × 10^-2 - 1 mol.m^-3; optimum ca. 1–2 × 10^-1mol.m^-3). At the optimum concentration of PAA, the stimulation of both responses was about 63–75% of that induced by the optimum concentration of IAA. The differences in the concentration range and magnitude of the responses to IAA and PAA were not due to differences in uptake of the two compounds. The stimulation of elongation by both compounds was prevented by 3.6 × 10^-2mol.m^-3 cycloheximide (CH), and acid invertase activites were greatly reduced compared with samples treated with growth substances alone. A saturating concentration of the specific auxin efflux carrier inhibitor N-1-naphthylphthalamic acid (NPA) slightly promoted the growth of control segments, probably by reducing the loss of residual endogenous auxin to the incubation medium. The elongation induced by PAA at its optimum concentration was considerably greater than the elongation induced by NPA, indicating that PAA did not cause growth by preventing the loss of endogenous auxin from the segments. Elongation responses to combinations of IAA and PAA suggested that the compounds were acting additively and that they were affecting growth by the same mechanism.     

Somatic mutations caused by excision of the transposable element, Tpn1, from the DFR gene for pigmentation in sub-epidermal layer of periclinally chimeric flowers of Japanese morning glory and their germinal transmission to their progeny

Pigmentation in flowers of Japanese morning glory is intense in the epidermal layer, lighter in the subepidermis, and much lighter in the internal tissues; by contrast coloration in stems occurs only in the sub-epidermal layer. The a-3 ^f mutant of Japanese morning glory bears white flowers with normal-colored flecks and sectors, and its variegation also occurs in leaves and stems. The mutable line can produce chimeric flowers pigmented uniformly in the sub-epidermal tissue and variegated in the epidermal layer, and stems of these flowers are also pigmented. Since they give selfed progeny that segregate to give a ratio of three germinal revertants bearing fully colored flowers to one flecked mutant, it has been [OR Imai (1934) has] postulated that somatic mutations in the sub-epidermal layer can be transmitted to the next generation and that the germ cells in the reproductive organs must form from the cells of the sub-epidermal layer. Recently, we found that the 6.4-kb En/Spm-related transposable element, Tpn1, resides within the DFR-B gene for anthocyanin biosynthesis in the mutable a-3 ^f line. To test whether somatic mutations caused by Tpn1 excision from the DFR-B gene in the subepidermis of periclinally chimeric flowers are transmissible to their progeny, we have examined the structure of the DFR-B region in the germinal revertants derived from the chimeric flowers and compared the sequences generated by the somatic excision of Tpn1 in periclinally chimeric flowers with those in their germinal revertants. Our results confirm that somatic mutations caused by Tpn1 excision from the DFR-B gene in the sub-epidermal tissue of chimeric flowers can be transmitted to their progeny, which results in the generation of germinal revertants.     

Temperature-dependent internode elongation in vegetative plants of Arabidopsis thaliana lacking phytochrome B and cryptochrome 1

 Vegetative plants of Arabidopsis thaliana (L.) Heynh. form a compact rosette of leaves in which internode growth is virtually arrested. Rapid extension of the internodes occurs after flower buds are present in the reproductive apex. Under natural radiation, continuous light from fluorescent lamps, or short photoperiods of light from fluorescent lamps, plants of the phyB cry1 double mutant (lacking both phytochrome B and cryptochrome 1) did not form normal rosettes because all the internodes showed some degree of elongation. Internode elongation was weak in the phyB single mutant and absent in the cry1 mutant, indicating redundancy between phytochrome B and cryptochrome 1. The absence of phytochrome A caused no effects. The failure to form normal rosettes was conditional because internode elongation was arrested at low temperatures in all the mutant combinations. In contrast, the temperature dependence of phytochrome B and cryptochrome 1 effects on hypocotyl growth was weak. The elongation of the internodes in phyB cry1 was not accompanied by early flowering as showed by the lack of effects on the final number of leaves. Apex dissection indicated that in phyB cry1 double mutants internode elongation anticipated the transition from the vegetative to the reproductive stage. Thus, stem growth in Arabidopsis thaliana is not fully dependent on the program of reproductive development. Received: 2 June 1999 / Accepted: 13 August 1999     

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.     

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.     

The role of nodal regions in growth of Xanthium (Compositae) stem

Vegetative Xanthium plants grown under noninductive conditions were marked with India ink along the stem and photographed at three consecutive time intervals. Relative elemental rates (d[dX/dt]/dX) of stem elongation were estimated from displacement of marks during stem elongation. Young internodes elongated with constant relative elemental rates of 0.2 day^-1. Older internodes displayed an acropetal pattern of elongation in which the basal segments of an internode stopped elongating first and the apical portion last. Nodal regions elongated with very small relative elemental rates of 0.05 day^-1. Rates decreased as the age of the internodes and nodes increased and stopped shortly after Leaf Plastochron Index 9.     

Gibberellin Effect on Tryptophan Metabolism, Auxin Destruction, and Abscission in Coleus

Application of gibberellic acid (GA) to the apical region of the stem enhances ¹⁴CO₂ release from tryptophan-l-¹⁴C in cell free preparations of the apical region. Although GA when applied to the apical region markedly accelerates abscission rates of debladed petioles at the 4th node, the enhancement effect on tryptophan metabolism appears to be restricted to the apical bud region. The increased levels of diffusible auxin in Coleus stems, observed earlier by Muir and Valdovinos (1965), appear to be due to the GA effect on auxin precursor conversion rather than to an altered rate of auxin destruction. GA pre-treatment does not significantly alter destruction rates of auxin in the stem tissue. This is demonstrated by the release of ¹⁴CO₂ from IAA-1-¹⁴C by sections of internode tissue. While a multiple deblading pattern retards abscission of debladed petioles considerably, application of GA to debladed petioles at the basal region of the stem restores the normal rates of abscission at debladed distal nodes. No significant change in the abscission rates at treated nodes is observed. The GA effect on abscission at distal nodes is attributed to the effect of the growth substance on auxin precursor conversion in the apical region. In these experiments, as in the case of plants treated in the apical region with GA, auxin destruction rates in the stem are not altered significantly.