Propagation of Hedera helix: Influence of irradiance to stock plants, length of internode and topophysis of cutting

Abstract Stock plants of Hedera helix cv. Pittsburgh were grown in controlled environment rooms at four different irradiances (light intensities) 10, 22, 37 or 46 W m^-2 PAR (photosynthetically active radiation). The root formation on single internode cuttings from these stock plants was observed in relation to length of the internodes and the position on the vine (topophysis). The analysis indicated that the root number was primarily dependent on internode length which in turn was dependent on irradiance to stock plants and topophysis. The irradiance of 37 W m^-2 PAR was optimal for internode length and root number and the basal internode was the one which produced the greatest number of roots.     

Uncoupling light quality from light irradiance effects in Helianthus annuus shoots: putative roles for plant hormones in leaf and internode growth

An attempt has been made to uncouple the effects of the two primary components of shade light, a reduced red to far-red (R/FR) ratio and low photosynthetically active radiation (PAR), on the elongation of the youngest internode of sunflower (Helianthus annuus) seedlings. Maximal internode growth (length and biomass) was induced by a shade light having a reduced R/FR ratio (0.85) under the low PAR of 157 micromol m(-2) s(-1). Reducing the R/FR ratio under normal PAR (421 micromol m(-2) s(-1)) gave similar growth trends, albeit with a reduced magnitude of the response. Leaf area growth showed a rather different pattern, with maximal growth occurring at the higher (normal) PAR of 421 micromol m(-2) s(-1)), but with variable effects being seen with changes in light quality. Reducing the R/FR ratio (by enrichment with FR) gave significant increases in gibberellin A(1) (GA(1)) and indole-3-acetic acid (IAA) contents in both internodes and leaves. By contrast, a lower PAR irradiance had no significant effect on GA(1) and IAA levels in internodes or leaves, but did increase the levels of other GAs, including two precursors of GA(1). Interestingly, both leaf and internode hormone content (GAs, IAA) are positively and significantly correlated with growth of the internode, as are leaf levels of abscisic acid (ABA). However, changes in these three hormones bear little relationship to leaf growth. By implication, then, the leaf may be the major source of GAs and IAA, at least, for the rapidly elongating internode. Several other hormones were also assessed in leaves for plants grown under varying R/FR ratios and PARs. Leaf ethylene production was not influenced by changes in R/FR ratio, but was significantly reduced under the normal (higher) PAR, the irradiance treatment which increased leaf growth. Levels of the growth-active free base and riboside cytokinins were significantly increased in leaves under a reduced R/FR ratio, but only at the higher (normal) PAR irradiance; other light quality treatments evoked no significant changes. Taken in toto, these results indicate that both components of shade light can influence the levels of a wide range of endogenous hormones in internodes and leaves while evoking increased internode elongation and biomass accumulation. However, it is light quality changes (FR enrichment) which are most closely tied to increased hormone content, and especially with increased GA and IAA levels. Finally, the increases seen in internode and leaf GA content with a reduced R/FR ratio are consistent with FR enrichment inducing an overall increase in sunflower seedling GA biosynthesis.     

REGULATORY ROLE OF INDOLE-3-ACETIC ACID AND GIBBERELLIC ACID IN VEGETATIVE DEVELOPMENT OF XANTHIUM PENNSYLVANICUM

A single application of gibberellic acid to young internodes significantly accelerated the rate of internode growth and the rate of leaf production in shoots of Xanthium pennsylvanicum Wallr. The average duration of one plastochron in treated plants was reduced to 43% of control levels. Gibberellic acid also had a pronounced morphogenetic effect on leaves so that the area and leaf length of treated plants were both significantly reduced. Depending upon concentration, auxin had both inhibitory and promotive effects on Xanthium shoots. Indole-3-acetic acid markedly altered the response of the gibberellic acid-treated internodes and those located above and below the site of application. In addition, high auxin concentrations induced the formation of adventitious roots in treated internodes. Auxin also brought about significant reductions in the length and area of leaves developed under the influence of this hormone.     

光照强度和基质养分对匍匐茎草本金戴戴克隆生长和克隆形态的影响

在深度遮光(光照强度为高光条件的6.25%,约为自然光照的5.3%)或低养分条件下,金戴戴(Halerpestes ruthenica Ovcz.)生物量、初级分株叶面积、分株总数、匍匐茎总数和总长度均显著减小,而比节间长和比叶柄长显著增加.在低养分条件下,金戴戴匍匐茎平均节间长显著增加,而匍匐茎分枝强度和分株数显著减小.这些结果与克隆植物觅食模型相符合,表明当生长于异质性生境中,金戴戴可能通过以克隆生长和克隆形态的可塑性实现的觅养行为来增加对养分资源的摄取.在深度遮光条件下,金戴戴平均间隔子长度(即平均节间长和平均叶柄长)均显著减小.这一结果与以往实验中匍匐茎草本间隔子对中度和轻度遮光(光照强度为高光条件的13%～75%,>10%的自然光照)的反应不同.这表明,在深度遮光条件下匍匐茎克隆植物可能不发生通过间隔子可塑性实现的觅光行为.光照强度和基质养分条件的交互作用对许多性状如总生物量、匍匐茎总数和总长度、二级和三级分株数、分株总数、初级分株叶面积以及分枝强度均有十分显著的效应.在高光条件下,基质养分对这些性状有十分显著的影响;而在低光条件下,基质养分条件对这些性状不产生影响或影响较小.这表明,光照强度影响金戴戴对基质养分的可塑性反应.在深度遮光或低养分条件下,金戴戴可能通过减小匍匐茎节间粗度(增加比节间长)来增加或维持其相对长度,从而更有机会逃离资源丰度低的斑块.     

光照强度和基质养分对匍匐茎草本金戴戴克隆生长和克隆形态的影响

在深度遮光 (光照强度为高光条件的 6 .2 5% ,约为自然光照的 5.3% )或低养分条件下 ,金戴戴 (HalerpestesruthenicaOvcz.)生物量、初级分株叶面积、分株总数、匍匐茎总数和总长度均显著减小 ,而比节间长和比叶柄长显著增加。在低养分条件下 ,金戴戴匍匐茎平均节间长显著增加 ,而匍匐茎分枝强度和分株数显著减小。这些结果与克隆植物觅食模型相符合 ,表明当生长于异质性生境中 ,金戴戴可能通过以克隆生长和克隆形态的可塑性实现的觅养行为来增加对养分资源的摄取。在深度遮光条件下 ,金戴戴平均间隔子长度 (即平均节间长和平均叶柄长 )均显著减小。这一结果与以往实验中匍匐茎草本间隔子对中度和轻度遮光 (光照强度为高光条件的 1 3%～ 75% ,>1 0 %的自然光照 )的反应不同。这表明 ,在深度遮光条件下匍匐茎克隆植物可能不发生通过间隔子可塑性实现的觅光行为。光照强度和基质养分条件的交互作用对许多性状如总生物量、匍匐茎总数和总长度、二级和三级分株数、分株总数、初级分株叶面积以及分枝强度均有十分显著的效应。在高光条件下 ,基质养分对这些性状有十分显著的影响 ;而在低光条件下 ,基质养分条件对这些性状不产生影响或影响较小。这表明 ,光照强度影响金戴戴对基质养分的可塑性反应。在深度遮光     

Direct shoot regeneration from leaf and internode explants of Aloysia polystachya [GRIS.] mold. (Verbenaceae)

Summary Adventitious bud regeneration from leaf and internode explants of Aloysia polystachya was achieved. Shoots from nodal segments grown in vitro were cut into pieces and used as sources of explants. Organogenesis was induced from both explants cultured on quarter-strength Murashige and Skoog (MS) semisolid medium (plus sucrose 5 g l⁻¹) containing different combinations of 6-benzyladenine (BA) and α-naphthaleneacetic acid (NAA) under 116 μmol m⁻² s⁻¹ photosynthetic photon flux density (PPFD), 14-h photoperiod, and at a temperature of 27±2°C. The type of explant markedly influenced organogenesis and growth of the regenerated shoots. The regeneration frequencies were higher with leaf explants, while the number of shoots formed per responsive explant was greater with internode explants. However, the growth of regenerated shoots from internodes was seriously affected by vitrification. The number of shoots produced per responsive leaf explant increased from one to seven as the percentage of leaf explants producing shoots increased from 20 to more than 80%. NAA at 0.05 μM in combination with BA at 0.5μM induced the highest regeneration rate (87±8.8%) after 20 d of culture, yielding 5.9±0.8 shoots per responsive leaf explant. Histological examination confirmed the occurrence of direct organogenesis. The regenerated shoots from the best induction treatment were transferred to a fresh medium without plant growth regulators for 30 d. Finally, the elongated shoots were rooted by pre-treatment in an aqueous solution of NAA at 500 μM for 2 h and transferred to 1/4 MS. All plantlets raised in vitro were phenotypically normal and successfully hardened to ex vitro conditions. An experimental field plot with 2-yr-old in vitro-regenerated plants was established.     

The inhibition of potato sprout growth by light.

The effect of light intensity on sprout growth in seed potato tubers (Solanum tuberosum) was examined using diffuse daylight in Peru and diffuse artificial light at Glasgow. Mean temperatures below 20 °C produced strong sprout growth that was inhibited by both daylight and artificial light, at visible irradiances above 0.01 Wm^-2. The percentage inhibition of sprout growth increased linearly with the logarithm of the irradiance, 50% inhibition being at 0.04 - 0.1 Wm^-2 provided that the temperature was suitable for substantial sprout growth in the absence of light. Cultivar and temperature had very little effect on the 50% inhibition point. At high irradiances growth inhibition was up to 95%, but the sprout length was never reduced to zero; short, robust green sprouts remained. Sprout numbers were increased by daylight, but not by artificial light. Diffuse daylight also reduced the total weight loss from seed tubers during a storage season of 180 days. At mean temperatures above 20 °C., sprout growth in the absence of light was much reduced and the effect of light on sprout elongation was less obvious.     

Correlative Growth in Young Terminalia superba in a Controlled Environment: Effect of the Leaves on Internode Elongation

Young Terminalia superba plants were cultivated in a controlledenvironment at the Phytotron. Effects of the excision of a youngleaf at definite elongation stages and at two given levels ofthe main axis were studied on the elongation of internodes inthese plants. Effects of the leaf did not seem to depend onits nodal position on the main axis but predominantly occurredon the immediate surrounding internodes. The excision of a youngleaf enhanced the growth rate of the internode located belowit and markedly decreased the elongation of the internode aboveit but slightly affected internode growth duration. This excisionenhanced the final length of the internode located below theleaf and decreased the final length of the internode locatedabove the removed leaf. Significant linear regressions werefound between the length of the excised leaf and the internodefinal lengths. Microscopic examination of epidermal cells ofcontrol and disrupted internodes revealed that the decreasedelongation after leaf excision could be attributed to reductionof cell divisions. The increased elongation after leaf excisioncould be attributed both to slight increase in the length ofcells (significant negative correlation was found with the lengthof the removed leaf) and to increase of cell divisions. Terminalia superba, leaf excision, cell division, internode elongation, correlative growth     

Development of shoot inHydrangea macrophylla I. Terminal and axillary buds

The structure of shoots, in particular of winter buds, ofHydrangea macrophylla was examined. The non-flower-bearing shoot is usually composed of a lower and an upper part, between which a boundary is discernible by means of a distinctly short internode. This internode is the lowermost of the upper part, and it is usually shorter than the internodes immediately above and below, although the internodes tend to shorten successively from the proximal to the distal part of the shoot. Variations exist in the following characters among the terminal bud, the axillary bud on the lower part of the shoot and the axillary bud on the upper part: (1) length of bud; (2) character of the outermost pair of leaf primordia; (3) degree of development of secondary buds in the winter bud; and (4) the number of leaf primordia. Usually, the terminal bud contains several pairs of foliage leaf primordia with a primordial inflorescence at the terminal of the bud, but the axiallary bud contains only the primordia of foliage leaves in addition to a pair of bud scales.     

Gibberellins in Diffusates from Shoots of Apple Trees

Excised shoot apices, leaves and internodes from shoots of apple trees (Malus×domestica) give off gibberellins by diffusion on agar. A methanol extract of the agar was prepared, the extract separated on thin layer plates, and the gibberellin activity estimated by means of Rumex and lettuce hypocotyl bioassays. The largest amounts of gibberellin are found in diffusates from the shoot apex, the two upper leaves and the two upper internodes. Several promotive fractions are found on the chromatograms as well as growth inhibitors. Removal of young leaves retards elongation of the internodes. Probably gibberellins produced in young leaves exercise some control over this process. The growth regulators Alar and CCC also retard internode elongation. Diffusates from shoots treated with these substances were also examined. Preliminary results suggest that the amount of diffusible gibberellins from treated shoots is not reduced.     

Internodes of grain legumes — New location for xylem parenchyma transfer cells

Summary Xylem parenchyma transfer cells were observed in the primary and secondary vascular tissue of stem internodes of 21 in 28 species of grain legumes. Their structural features were similar to those of other transfer cells. The relationships of these cells to transfer cells at nodes were investigated. Non-nodulated seedlings ofPhaseolus vulgaris L. formed internode transfer cells if provided mineral nutrients through their roots, but not if grown in distilled water or fed nutrients entirely through their leaves. Wall ingrowths formed in parenchyma of primary xylem ofPhaseolus just before full extension of an internode. The significance of this new location for transfer cells was discussed.     

Interrelationships between Light Intensity and the Physiological Effects of 2:4-Dichlorophenoxyacetic Acid on the Growth of Helianthus annuus

The interrelationships between light intensity and the effectsof sodium 2:4-dichlorophenoxyacetate on the growth of Helianthusannuus in the early vegetative stage have been studied by subjectingtreated and control plants to a range of light intensities from,1.0 to 0.12 daylight. The growth regulator in varying amountswas applied either as droplets of aqueous solution to the firstor second pairs of leaves or as an overall spray. When the amountof the compound is such as to cause small but significant reductionsin the relative growth rate, the leaf-area ratio and the ratioof leaf area to leaf weight are likewise depressed but the netassimilation rate is relatively unaffected. Between 0.25 daylightand full daylight the proportionate changes in the relativegrowth rate, leaf-area ratio, and net assimilation rate inducedby sub-lethal amounts of the growth regulator are not greatlymodified by the level of light. If the intensity is reducedfurther to 0.12 daylight, then the reactions of the shaded plantsdiffer markedly from those of unshaded plants. For example,when the plants are shaded both before and after a spray application,the concentration required to cause a 50 per cent. mortalityis one-tenth of that demanded for plants receiving full daylight.For less phytotoxic amounts the percentage reductions, relativeto the controls, induced in the growth rate of the shoot aremore dependent on the intensity after than before the applicationof either a spray or measured droplets. These greater depressions in shoot growth at 0.12 daylight afterthe application are linked with comparable depressions in therate of growth of the leaves of which the first pair are moresensitive. Similarly, the growth rate of the first internodeis also depressed more under shade conditions, but that of thesecond internode at low doses may be greatly increased. Forboth the leaves and internodes shading before as against shadingafter the application may have different significant effectson the changes in growth caused by the growth regulator. Undersome conditions the interactions between the light intensitybefore and after the application and the quantity of the growthregulator are exceedingly complex. By using sodium 2:4-dichloro-5-iodophenoxyacetatelabelled with radioactive I¹³¹, it has also been establishedthat the rate of penetration into leaves is accelerated by ahigh light intensity after the application, but that the intensityreceived prior to the application causes no significant effect.Evidence was also obtained that transport to the shoot fromthe treated leaf is more dependent on the light intensity afterthe application—the transport being greater in full daylight.It is concluded that a number of factors must be involved inbringing about these differential effects and their relativeimportance is discussed.     

How internode length,position and presence of leaves affect survival and growth of <Emphasis Type="Italic">Alternanthera philoxeroides</Emphasis> after fragmentation?

Disturbance is common in nature and disturbance-caused fragmentation of clones happens frequently in stoloniferous plants. After fragmentation storage in stolon internodes and leaves may enhance survival and growth of stoloniferous plants. We hypothesize that (1) increasing length of the internode attached to the ramet and (2) presence of leaves will increase ramet survival and growth, and that (3) internode positions (before or after the ramet or both) will also play a role. We tested these hypotheses with the stoloniferous, invasive herb Alternanthera philoxeroides. In one experiment, we measured survival and growth of the ramets either without stolon internode (0 cm in length) or attached with internodes of 2, 4, 6 and 8 cm and either with or without leaves. In the other experiment, we measured survival and growth of the ramets attached with a proximal internode (before the ramet), a distal internode (after the ramet) or both. Increasing internode length and presence of leaves significantly increased the survival rate and growth (biomass, leaf area, number of ramets, stolon length and number of leaves) of the A. philoxeroides plants. All growth measures of A. philoxeroides at harvest were larger when the ramets were attached with a distal internode than when they were attached with a proximal internode, but the survival rate was lower. These results support the hypotheses and suggest that storage in stolons and leaves may be of great significance for clonal plants in frequently disturbed habitats and may contribute greatly to the invasiveness of A. philoxeroides.     

Auxin Relations of the Woody Shoot: The Distribution of Diffusible Auxin in Shoots of Apple and Plum Rootstock Varieties

Surveys have been made of diffusible auxin in the stem tissuesof growing shoots of apple and plum rootstock varieties. Usingagar plates as carriers auxin was collected from the lower surfaceof isolated stem sections and assayed by the Avena curvaturemethod. The stool and layer shoots studied grew for severalmonths producing many leaves and reaching considerable lengths.The data provide information on selected internodes and showthe auxin status of the shoot at various times during growth,and the auxin gradients down the stem at these various times.Free auxin content of the shoot apex was consistently less thanthat of the internodes below. In 1946 auxin content declinedthroughout growth with a steady positioning of the auxin peakin the upper shoot. In 1947, following a period of drought,when growth almost ceased, a secondary auxin peak occurred positionedin lower internodes distant from the apex. This seasonal contrastwas reflected in the auxin relations of the individual internode,and was observed both in apple and plum. The nature of the auxindecline below the peak region, and the total disappearance offree auxin from the shoot as growth subsides, is discussed.The reappearance of free auxin in mature internodes, which hasnot been transmitted from the stem apex, implies either derivationfrom a stored state or the ability of the internode to produceits own auxin.     

The role of the leaves in the regulation of internode elongation in Phaseolus vulgaris

Time-course patterns of leaf and internode elongation were studied in bean plants. Each leaf started its main elongation period when the leaf below reached half of its final length. The onset of leaf unfolding was nearly synchronous with the initiation of the elongation of the subjacent internode. Excision of young leaves increased the rate of stem elongation as a result of an earlier unfolding of the next upper leaves and the concomitant advancement in the elongation of their subjacent internodes. IAA or NAA (1% in lanolin) suppressed the enhancement effects of leaf excision on leaf and internode elongation. The excision of a young leaf increased the final length of internodes located below it, and at the same time decreased the final length of the internodes located above the excised leaf. The reduction was greater the younger the internode. Differences in internode elongation after leaf excision were related to changes during internode ontogenesis in their relative response to the availability of assimilates on the one hand, and on the other hand to hormonal factors transported acropetally from the young leaves to the growing internodes.     

Ectopic expression of class 1 KNOX genes induce adventitious shoot regeneration and alter growth and development of tobacco (Nicotiana tabacum L) and European plum (Prunus domestica L)

Transgenic plants of tobacco (Nicotiana tabacum L) and European plum (Prunus domestica L) were produced by transforming with the apple class 1 KNOX genes (MdKN1 and MdKN2) or corn KNOX1 gene. Transgenic tobacco plants were regenerated in vitro from transformed leaf discs cultured in a medium lacking cytokinin. Ectopic expression of KNOX genes retarded shoot growth by suppressing elongation of internodes in transgenic tobacco plants. Expression of each of the three KNOX1 genes induced malformation and extensive lobbing in tobacco leaves. In situ regeneration of adventitious shoots was observed from leaves and roots of transgenic tobacco plants expressing each of the three KNOX genes. In vitro culture of leaf explants and internode sections excised from in vitro grown MdKN1 expressing tobacco shoots regenerated adventitious shoots on MS (Murashige and Skoog 1962) basal medium in the absence of exogenous cytokinin. Transgenic plum plants that expressed the MdKN2 or corn KNOX1 gene grew normally but MdKN1 caused a significant reduction in plant height, leaf shape and size and produced malformed curly leaves. A high frequency of adventitious shoot regeneration (96%) was observed in cultures of leaf explants excised from corn KNOX1-expressing transgenic plum shoots. In contrast to KNOX1-expressing tobacco, leaf and internode explants of corn KNOX1-expressing plum required synthetic cytokinin (thidiazuron) in the culture medium to induce adventitious shoot regeneration. The induction of high-frequency regeneration of adventitious shoots in vitro from leaves and stem internodal sections of plum through the ectopic expression of a KNOX1 gene is the first such report for a woody perennial fruit trees.     

Differential response to shading in orthotropic and plagiotropic shoots of the clonal herb Glechoma hirsuta

Previous studies have shown that internodes and petioles of closely related erect and stoloniferous species show marked differences in their response to shading. Vertical structures show strong elongation responses while horizontal structures show significantly smaller elongation responses. This paper reports an experiment designed to test whether internodes and petioles on orthotropic (vertically oriented) and plagiotropic (horizontally oriented) shoots produced by the same plant, also show such differential responses. The study species, Glechoma hirsuta, produces plagiotropic shoots in its vegetative stage and orthotropic shoots during its generative stage. Shoots of G. hirsuta were grown either in full daylight or under simulated canopy shade. Internode and petiole elongation, biomass investment patterns and other growth-related parameters were measured on plants in each light treatment. In orthotropic shoots the length of internodes responded more strongly to shading than the length of petioles, while the opposite was true for plagiotropic shoots, confirming the hypothesis that vertical spacers have higher degrees of shade-induced plasticity than horizontally oriented spacers. Growth and development of horizontally oriented shoots was reduced by shading, whereas that of vertically oriented shoots was unaffected by light treatment. The results are discussed in terms of the differences in function of vertically and horizontally oriented spacers, and the probable benefits of plasticity in enhancing essential activities including photosynthesis, habitat exploration and seed dispersal. Received: 23 December 1996 / Accepted: 19 July 1997     

Gibberellic acid improved shoot multiplication in Cephaelis ipecacuanha

The internodes of Cephaelis ipecacuanha elongated when cultured on Gamborg B5 solid medium supplemented with 0.5 or 1 mg/L gibberellic acid (GA₃). The size of the elongated internode doubled in length compared to the untreated shoots, and the adventitious shoots formed on the elongated internodes. The shoots grew easily into plantlets without the use of auxin for rooting. The ex vitro regenerates cultivated in the greenhouse showed normal characteristics. Emetic alkaloids were detected in the leaves of in vitro shoots and the roots of regenerates cultivated in the greenhouse. This method using GA₃ propagated numerous plants at a rate of more than 100 times compared to the method without GA₃.     

Implications of opposite phyllotaxis for light interception efficiency of Mediterranean woody plants

Opposite leaves lead to a greater leaf overlapping than leaves spirally arranged along a shoot, decreasing light interception efficiency (Ea, fraction of the light reaching the plant actually intercepted by the leaves) of the crown. However, Ea results from a whole suite of morphological traits. The interplay between phyllotaxis, crown architecture, leaf morphology and Ea was explored in 12 woody species from Mediterranean-type ecosystems, where the abundance of woody species with opposite phyllotaxis is unusually high. The three-dimensional model Y-plant was used to estimate Ea in unbranched, vertical shoots of each species encompassing the natural morphological variation found from moderate shade to open light environments. Ea exhibited significant interspecific differences, ranging from 0.25 in Daphne gnidium to 0.75 in Cistus ladanifer, Olea europaea and Salvia officinalis, decreasing with leaf inclination angle and leaf area ratio (LAR), and increasing with internode-to-leaf-length ratio and supporting biomass. Species with spiral vs. opposite phyllotaxis did not differ in their mean Ea. However, the former had higher Ea than the latter at short internode lengths. The natural range of variation in internode length had a larger effect on Ea than the natural range of leaf elevation angle. Principal component analysis segregated species with opposite phyllotaxis from those with spiral leaves because of their greater self-shading for high sun elevation angles (>45°); they were in turn distributed in two groups, one with high Ea, large investment in supporting biomass and long internodes, and another with low Ea and large LAR. Species with spiral phyllotaxis all had intermediate or low Ea and steep leaf elevation angles. Species with opposite phyllotaxis can compensate their less efficient leaf arrangement by decreasing leaf elevation angle and increasing internode length, but they may experience a real phylogenetic constraint for light interception when biomass allocation to supporting tissues (internodes and petioles) becomes very costly. This constraint could be involved in the shade intolerance of woody Mediterranean species exhibiting opposite phyllotaxis.