A Study on the Mechanism of Nodding Initiation of the Flower Stalk in a Poppy, Papaver Rhoeas L.

A study was made to find whether the nodding of the flower stalkin a poppy, Papaver Rhoeas L., immediately after its formationwas triggered by the weight of its flower bud or by positivegeoreaction and the following results were obtained.

1. The direction of the nodding was mostly toward the inclinedside of the stalk, which was opposite the leaf, for apical flowerbuds.
2. If the weight of the flower bud at stage 1 was cancelledbyapplying a load equivalent to the bud weight, the noddingofthe stalk was not initiated.
3. The stalk at stages 1 and2 and the upper part of the stalk(bending zone), as comparedwith the basal part, at later stageswere highly deformableaccording to measurements by the bendingmethod.
4. The cellwall of highly deformable stalks was rich in hemicellulosesand that of the basal part was abundant in pectic substances.

From these results, we concluded that the initiation of thenodding in the flower stalk was caused by the weight of theflower bud and positive geotropic reaction was probably notinvolved. (Received December 22, 1980; Accepted January 26, 1981)     

Hydrogen ion-induced curvature in cucumber hypocotyls

A strip of filter paper saturated with a pH 3.0 buffer solutionand placed on the upper side of a light-grown cucumber hypocotylseedling can eliminate the negative geotropic curvature of horizontallyplaced seedlings, by promoting elongation of the upper side.These filter paper strips also caused strong curvature of verticallyplaced seedlings and induced cell wall loosening of the epidermis. (Received February 16, 1973; )     

Root geotropism and the role of growth regulators from the cap: a re-examination

Abstract. It is widely believed that the root cap participates in geotropism by acting both as a sensor of the direction of gravity and as a source of at least one regulator of root growth, which may be abscisic acid (ABA). It has been suggested that this regulator accumulates within the prospective concave (or lower) half of the root and there causes a retardation of growth that brings about bending. A re-examination of the evidence upon which this inhibitor hypothesis of geotropic control is based reveals that (1) it is derived almost entirely from microsurgical studies and the deductions from such experiments still require corro-orations from analyses of inhibitor content and action; (2) the evidence that ABA is the inhibitor seems poor at present; (3) in maize and lentil, two well-studied species, geocurvature is probably a consequence of accelerated growth within the prospective convex (or upper) half rather than inhibited growth within the concave (or lower) half; (4) the geotropic signal from the cap may be one that redirects a pre-existing basipetal flow of inhibitor away from the upper     

Changes in diffusible indole-3-acetic acid from various parts of tulip plant during rapid elongation of the flower stalk

In order to chemically identify the putative indole-3-acetic acid (IAA) and to confirm the native source of auxins account for rapid elongation of the floral stalk of tulip, we examined diffusible IAA from various parts of tulip plant during rapid elongation of the flower stalk. IAA was identified in the diffusates collected from the leaves, internodes, and floral organs with gas chromatography (GC)–mass spectrometry. The amount of diffusible IAA from different plant organs followed the order of that the internodes > flower organs > leaves during the period of rapid elongation of the floral stalk. The diffusible IAA from internodes reached its peak amount at different time than did diffusible IAA from the flower. The results obtained indicated that the top internode is probably the major source of auxins account for rapid elongation of the flower stalk.     

The Diageotropic Behaviour of Rhizomes

1. The geotropic behaviour of rhizomes of Aegopodium podagrariahas been investigated by time-lapse photographic recording usinginfra-red radiation in complete absence of light.
2. They havebeen shown to be very sensitive to light. Even a singleexposurefor 30 seconds to red light evokes a characteristicresponsein which the rhizome first turns downwards and thenturns upagain to the horizontal position. The exposure to lightduringthe process of digging up the rhizomes and setting upin theapparatus has similar effects which die away after about24hours.
3. After recovery from this initial light effect theycontinueto grow roughly horizontal if maintained in darknessor subjectedto infra-red radiation only. If inverted (turnedthrough 180?)the side which was lowermost during the inversiongrows forabout 2 hours relatively the more quickly, thus becomingconvex;then a rapid straightening and bending in the oppositedirectionoccurs. This is followed by further up-and-down movementswhichgradually die out and the rhizome then continues growingstraight.
4. Inversion for 10 minutes also evokes an essentiallysimilarresponse, which begins about 20 minutes after the invertedrhizomehas been returned to its original position. The importanttheoreticalsignificance of this is discussed in the text.
5. Displacementof rhizomes through angles other than 180? resultsin to-and-frobendings in which the side of the rhizome whichwas uppermostbefore the stimulus of displacement first becomesconvex. Therhizomes then turn gradually towards the horizontalplane.
6. Rhizomeswhich have recovered from the initial light effectand are thusgrowing in darkness horizontally are caused toturn upwardswhen the air surrounding them is replaced by air+5per cent,carbon dioxide.
7. The theoretical significance of these findingsis discussed.

     

Redistribution of Endogenous Indoleacetic Acid, Abscisic Acid and Gibberellins in Geotropically Stimulated Ribes nigrum Shoots

The presence of IAA, ABA and gibberellins in extracts of shoots of Ribes nigrum was demonstrated by gas-liquid chroma-tography (GLC) for both IAA and ABA and by the lettuce hypocotyl assay for gibberellins. Quantitative estimation of the three substances in extracts from upper and lower halves of shoots which had been kept horizontal, and which showed negative geotropic response after 4 h, indicated a redistribution of hormones during the geotropic stimulation. The ratio of the hormones in lower and upper halves was 3.8:1 and 2.8:1 for IAA and giberellins respectively, whereas the ratio of ABA in upper and lower halves was 2.1:1. There is, however, no evidence for the participation of gibberellins and ABA in the early development of negative geotropic curvatures, since shoots of intact Vicia faba seedlings treated with 100 mg/1 solution of GA₃, ABA and 10 mg/1 IAA for 30 min prior to geotropic stimulation, developed negative geotropic curvatures, although shoots pretreated with 50 and 100 mg/1 IAA did not develop curvature.     

Computer-based video digitizer analysis of surface extension in maize roots

We used a video digitizer system to measure surface extension and curvature in gravistimulated primary roots of maize (Zea mays L.). Downward curvature began about 25 +/- 7 min after gravistimulation and resulted from a combination of enhanced growth along the upper surface and reduced growth along the lower surface relative to growth in vertically oriented controls. The roots curved at a rate of 1.4 +/- 0.5 degrees min-1 but the pattern of curvature varied somewhat. In about 35% of the samples the roots curved steadily downward and the rate of curvature slowed as the root neared 90 degrees. A final angle of about 90 degrees was reached 110 +/- 35 min after the start of gravistimulation. In about 65% of the samples there was a period of backward curvature (partial reversal of curvature) during the response. In some cases (about 15% of those showing a period of reverse bending) this period of backward curvature occurred before the root reached 90 degrees. Following transient backward curvature, downward curvature resumed and the root approached a final angle of about 90 degrees. In about 65% of the roots showing a period of reverse curvature, the roots curved steadily past the vertical, reaching maximum curvature about 205 +/- 65 min after gravistimulation. The direction of curvature then reversed back toward the vertical. After one or two oscillations about the vertical the roots obtained a vertical orientation and the distribution of growth within the root tip became the same as that prior to gravistimulation. The period of transient backward curvature coincided with and was evidently caused by enhancement of growth along the concave and inhibition of growth along the convex side of the curve, a pattern opposite to that prevailing in the earlier stages of downward curvature. There were periods during the gravitropic response when the normally unimodal growth-rate distribution within the elongation zone became bimodal with two peaks of rapid elongation separated by a region of reduced elongation rate. This occurred at different times on the convex and concave sides of the graviresponding root. During the period of steady downward curvature the elongation zone along the convex side extended farther toward the tip than in the vertical control. During the period of reduced rate of curvature, the zone of elongation extended farther toward the tip along the concave side of the root. The data show that the gravitropic response pattern varies with time and involves changes in localized elongation rates as well as changes in the length and position of the elongation zone. Models of root gravitropic curvature based on simple unimodal inhibition of growth along the lower side cannot account for these complex growth patterns.     

Comparative studies on auxin and fusicoccin actions on plant growth

Mode of action of FC was compared with that of auxin in differentexperimental systems and the following results were obtained.

1. FC, as well as auxin, primarily induced elongation of the epidermisof pea epicotyl segments, but it also promoted elongation ofthe inner tissue, as judged by its action in split stem tests,elongation of hollow-cylinder segments and elongation of unpeeledand peeled segments.
2. FC decreased the minimum stress relaxationtime (T₀) and increasedthe extensibility (mm/gr) of the epidermalcell wall of peaepicotyl segments, as did auxin.
3. FC failedto induce expansion growth of Jerusalem artichoketuber sliceswhen given alone or in combination with kinetinor gibberellicacid.
4. FC at concentrations lower than 10^–6 M, when givenwithauxin at concentrations lower than 0.03 mg/liter, promotedelongationof Avena coleoptile segments in an additive manner,to achievethe maximum elongation at higher concentrations.
5. An antiauxin, 2,4,6-trichlorophenoxyacetic acid, inhibitedtheelongation of Avena coleoptile segments due to auxin butnotthat due to FC.
6. Nojirimycin, an inhibitor of ß-glycosidases,inhibitedelongation of pea internode segments due not onlyto auxin butalso to FC.
7. At concentrations more than 10^–5MFC promoted root elongationof intact lettuce seedlings, whichwas inhibited by exogenousauxin.

From these results it is concluded that FC and auxin have acommon mechanism, which may involve hydrogen ion extrusion,leading to cell wall loosening and thus cell elongation. Thisgrowth is limited to the extent that the cells are capable ofelongating in response to hydrogen ions. Otherwise there isa definite difference in the mode of actions between FC andauxin, including the nature of cellular receptors for thesetwo compounds. (Received August 29, 1974; )     

Changes in the shape of epithelial embryonic cells of the spur-toed frog upon deformation of the cell layer

A quantitative study was made of changes in the shape of cells in double explants of the blastocoel roof of the clawed frog gastrula within the first four hours after artificial bending of explants. It was found that, on the concave (contracted) side of explants, epithelial cells stretched out, and in many of them the apical surface contracted, whereas on the convex (stretched) side the cells remained isodiametric. The maximal difference in the apical index between epithelial cells located on the concave and convex sides was observed after 2 h of explant cultivation; by 2 h the artificially produced curvature of the explant further increased. Endocytosis on the concave side was more active than on the convex side. Experiments with inhibitors modulating the behavior of the actomyosin complex showed that unimpeded functioning of myosin II is more important for the apical contraction and elongation of cells than proper structural organization of the actin backbone.     

Cell elongation in the grass pulvinus in response to geotropic stimulation and auxin application

Summary Horizontally-placed segments of Avena sativa L. shoots show a negative geotropic response after a period of 30 min. This response is based on cell elongation on the lower side of the leaf-sheath base (pulvinus). Triticum aestivum L., Hordeum vulgare L. and Secale cereale L. also show geotropic responses that are similar to those in Avena shoots. The pulvinus is a highly specialized organ with radial symmetry and is made up of epidermal, vascular, parenchymatous and collenchymatous tissues. Statoliths, which are confined to parenchyma cells around the vascular bundles, sediment towards the gravitational field within 10–15 min of geotropic stimulation. Collenchymatous cells occur as prominent bundle caps, and in Avena, they occupy about 30% of the volume of the pulvinus. Geotropic stimulation causes a 3- to 5-fold increase in the length of the cells on the side nearest to the center of the gravitational field. Growth can also be initiated in vertically-held pulvini by the application of indole-3-acetic acid, 1-naphthaleneacetic acid or 2.4-dichlorophenoxyacetic acid. 2.3.5.-triiodobenzoic acid interferes with growth response produced by geotropic stimulation as well as with the response caused by auxin application. Gibberellic acid and kinetin have no visible effect on the growth of the pulvinus. Polarization microscopy shows a unique, non-uniform stretching of the elongating collenchymatous cells. Nonelongated collenchymatous cells appear uniformally anisotropic. After geotropic stimulation or auxin application, they appear alternately anisotropic and almost isotropic. Such a pattern of cell elongation is also observed in collenchyma cells of geotropically-stimulated shoots of Rumex acetosa L., a dicotyledon.Abbreviations 2.4-D 2.4-dichlorophenoxyacetic acid - GA₃ gibberellic acid - IAA indole-3-acetic acid - NAA l-naphthaleneacetic acid - TIBA 2.3.5-triiodobenzoic acid     

Biological activity of phyllosinol, a phytotoxic compound isolated from a culture filtrate of Phyllosticta sp.

1. Phyllosinol is a phytotoxic metabolite of Phyllosticta sp. Thissubstance at 100 µg/ml produced dark grey necrotic lesionson the leaf of red clover. Sensitivities of various plant speciesto phyllosinol differed both quantitatively and qualitatively.
2. Phyllosinol reduced root growth in rice seedlings by 60% at10^–4 M, whereas stimulation of root elongation occurredat a concentration range from 10^–9 to 10^–5 M.
3. Phyllosinolat 2.5x10^–4M promoted adventitious root formationin epicotylsof Azukia cuttings by about 100%. Promotion waspartly reducedby simultaneous application of cysteine.
4. IAA-induced elongationof isolated Avena coleoptile sectionswas inhibited by phyllosinolat a concentration range from 10^–5to 10^–3M.
5. Sulfhydrylcompounds, i.e. cysteine and glutathione relievedinhibitioncaused by phyllosinol in IAA-induced elongation ofAvena coleoptilesections.
6. GA₃-induced elongation of wheat leaf sections wasslightly inhibitedby phyllosinol at 10^–4M.
7. Phyllosinolalso has antibiotic activity. Among the organismstested, Phycomycetesand Gram-negative bacteria appeared mostsusceptible to phyllosinol.

(Received April 21, 1970; )     

Stipe elongation during basidiocarp maturation in Coprinus macrorhizus: Mechanical properties of stipe cell wall

Stipe elongation during basidiocarp maturation in the wild-type,#5026+5132, and the elongationless mutant, NG0398, of Coprinusmacrorhizus was studied, and the following results were obtained.

1. In the wild-type the middle zone of the stipe elongated 8.4times in 15 hr during maturation, while in the mutant it elongatedoaiy 2.2 times.
2. Component cells of the stipe elongated inparallel with thestipe elongation in both the wild-type andthe mutant. The widthof stipe cells was almost constant duringelongation in thewild-type, while it increased 2 times in themutant. Cell volumeincreased ca. 8 times in both stocks.
3. Theosmotic value of stipe cells was almost constant (0.45–0.50M) throughout elongation of both the wild-type and the elongationlessstipes.
4. Mechanical properties of the cell wall were examinedby measuringshrinkage, extensibility and minimum stress-relaxationtime(To) of the stipe during maturation. These parameters weredirectlyproportional to the elongation rate to follow.
5. Whenthe wild-type stipes were incubated in various concentrationsof mannitol solution and then in plain buffer solution, theextensibility of the stipe after the incubation in mannitolsolutions changed proportionally with the stipe length afterthe mannitol treatment, and To with the elongation capacityin plain buffer solution.

(Received March 3, 1977; )     

Effects of gibberellic acid and indole-3-acetic acid on stress-relaxation properties of pea hook cell wall

The effects of GA, IAA and PCIB on the cell wall propertiesof Alaska pea hooks were examined using stress-relaxation analysis.The results were:

1. GA caused a decrease in the stress-relaxation parameter To ofplumular hook sections after the first 30 min of incubation,long before it induced elongation.
2. PCIB increased To, andIAA tended to negate the PCIB effecton To in GA-treated sectionsafter 90 min of incubation, whenthe effect of PCIB and IAAon the elongation was not yet found.In this case, IAA couldnot be substituted by an extra amountof GA.
3. GA decreasedTo in the middle part of the sections after 24hr of incubation,and then stimulated elongation.
4. In any case, the effect ofGA, IAA or PCIB on To was recognizedin both epidermis and innertissue of plumular hook sections.
5. The stress-relaxation parameterTo appears to represent thecapacity of the cell wall to extend;we thus concluded thatboth gibberellin and auxin increase theextensibility of thecell wall, when they stimulate the elongationof plumular hooksections.

(Received October 4, 1974; )     

Soluble and insoluble invertase activity in elongating Tulipa gesneriana flower stalks

Previously 'frozen' Tulipa gesneriana L. bulbs cv. Apeldoorn, were planted and grown at higher temperatures to study the role of invertase (EC 3.2.1.26) in the cold-induced elongation of the flower stalk internodes. After planting, flower stalks were left intact, or, the leaves and flower bud were both removed to inhibit internode elongation. In intact flower stalks, elongation of the internodes was accompanied by an accumulation of glucose and an initial decrease in the sucrose content g,⁻¹ dry weight. Insoluble invertase activity g,⁻¹ dry weight hardly changed, but soluble invertase activity showed a peak pattern, that was related, at least for the greater part, to the changes in the sugar contents. Peak activities of soluble invertase were found during (lower- and uppermost internodes) or around the onset of the rapid phase of internode elongation (middle internodes). Internode elongation and glucose accumulation immediately ceased when the leaves and flower bud were removed. Insoluble invertase activity g,⁻¹ dry weight remained at its initial level (lowermost internode) or increased more towards the upper internodes. Soluble invertase activity did not further increase (uppermost internode) or decreased abruptly to a low level. It is concluded that soluble invertase may be one of the factors contributing to glucose accumulation and internode elongation in the tulip flower stalk.     

Physiological Studies in the Mucorales: PART I THE PHOTOTROPISM OF SPORANGIOPHORES OF PHYCOMYCES BLAKESLEEANUS

Previous views on the physical basis of phototropism in Phycomycessporangiophores are briefly discussed.

1. It was confirmed thatunilaterally illuminated sporangiophoresimmersed in liquidparaffin show strong negative phototropism.
2. Elongation growthceased and no phototropic response took placeunder anaerobicconditions.
3. By focusing a fine beam of light on to one edgeof the growingzone of a sporangiophore, leaving the other sidein darkness,it was established that greater elongation tookplace in theilluminated zone, the sporangiophore tending tobend out ofthe beam. Rapid reversal of the curvature followedwhen theillumination was transferred to the opposite edge ofthe sporangiophore.

Wassink and Boumann's suggestion that phototropism can be initiatedby a one-quantum-per-cell process is criticized in the lightof this result and other work by Castle.     

FURTHER STUDIES ON THE ACTION OF FREE AMINO ACIDS ON FLOWERING OF DUCKWEED, LEMNA GIBBA G3

1. The maximum inhibition of flowering of a long-day duckweed,L. gibba G3, occurred when amino acid was added to the culturemedium on the third long-day. Lysine could reverse the inhibitionnon-specifically.
2. When the concentration ratio of endogenousarginine to endogenouslysine was under 10, the flowering processesprogressed normally,and the flower production decreased withthe increase of theratio from 10 to 20.
3. These and relevantfindings support our previous conclusionthat in vivo free aminoacids may contribute in establishingintracellular state determiningwhether certain essential reaction(s)involved inflowering processescan proceed normally or not.

( Accepted February 15, 1965)     

The Effects of Gibberellins on the Growth of Excised Tomato Roots

1. At appropriate concentrations both gibberellic acid (GA) and1-naphthalene-acetic acid (NAA) enhance the main axis growthof excised tomato roots grown in culture media containing sucroseat concentrations below 1 per cent. Lateral root extension growthis enhanced by GA at all sucrose concentrations tested; onlyat the lower sucrose concentrations is this effect observedwith NAA. Both GA and NAA increase the number of emergent lateralroots and this effect is most marked in media of low sucrosecontent. Both GA and NAA at higher concentrations inhibit rootgrowth but NAA exhibits its full range of growth effects overa much narrower concentration range than GA.
2. GA, like NAA,speeds up the loss of meristematic activity whichoccurs whenindividual meristems are repeatedly subculturedin media containing1 per cent, or higher concentrations ofsucrose.
3. The promotionof main axis growth by both GA and NAA involvesenhanced cellelongation and cell division. At a moderatelyinhibitory concentrationGA reduces both cell elongation andcell division; this is notthe case with NAA.
4. Gibberellins A₁, A₂, and A₄ resemble GA(gibberellin A₃) intheir growth effects. Allogibberic acidlike G A promotes lateralroot extension growth but causes markedinhibition of root growthat a much lower concentration thanGA.

     

GEOTROPIC CURVATURE OF AVENA COLEOPTILES

Applying a photographic recording method, and working on enlargements of the plates so obtained, the shape of the geotropically curving coleoptile of Avena was studied. This shape is expressed in terms of curvature of the neutral axis of the coleoptile; for the true geotropic response of the strain of Avena used, the curvature of the neutral axis is an arc of a circle. The "rate of curvature" is taken as the derivative of the curve relating time with the angle with the horizontal made by the tangent to the neutral axis at the tip. This rate increases up to a maximum and then decreases gradually. No "geogrowth" of the whole coleoptile is found. It is shown that the curvature is due to an increase in the elongation of the lower side of the horizontally placed coleoptile with a concomitant decrease of the rate of elongation of the upper side. This is correlated with a shift in distribution of "growth substance" in the tip as affected by change of position of the coleoptile.     

Distribution of labeled auxin in geotropically stimulated steins of cucumber and pea

The distribution of IAA-2-¹⁴C or IAA-5-³H applied to the apexin the upper and lower (with respect to gravity) halves of geotropicallystimulated stems of cucumber and pea was examined and the resultsobtained are as follows: 1. A larger amount of IAA-2-¹⁴C or IAA-5-³H was detected inthe lower than upper half of cucumber hypocotyls with 3-hr geostimulation. 2. A larger amount of IAA-2-¹⁴C was distributed in the lowerthan upper epidermis of pea epicotyls with 1-hr geostimulation. 3. Freezing autoradiography revealed that IAA-2-¹⁴C was concentratedin the vascular bundles and epidermis of cucumber hypocotyls,the distribution being affected by geostimulation only in thelatter. 4. Application of 1% TIBA in lanolin inhibited the distributionof IAA-2-¹⁴C in the lower epidermis of pea epicotyls, causingsuppression of geotropic curvature. 5. From these results, we concluded that IAA which accumulatedin the lower epidermis of the stem upon geostimulation causedthe negative geotropic curvature of the stem. (Received October 13, 1975; )     

The source and lateral transport of growth inhibitors in geotropically stimulated roots of Zea mays and Pisum sativum

Summary The positive geotropic responses of the primary roots of Zea mays and Pisum sativum seedlings depend upon at least one growth inhibiting factor which arises in the root cap and which moves basipetally through the apex into the extending zone. The root apex (as distinct from the cap) and the regions more basal to the extending zone are not sources of growth regulators directly involved in the geotropic response. A difference in the concentration or effectiveness of the inhibitory factor(s) arising in the cap must be established between the upper and lower halves of a horizontal root. Positive geotropic curvature in a horizontal root is attributable, at least in part, to a downward lateral transport of inhibitor(s) from the upper to the lower half of the organ.