Effects of Mecoprop (an Auxin Analogue) on Ethylene Evolution and Epinasty in Two Biotypes of Stellaria media

Petiolar epinasty and the production of ethylene (ethene) werestudied in chickweed biotypes, Stellaria media, treated withthe herbicide and auxin analogue (RS)-2-(4-chloro-o-tolyloxy)propionicacid, potassium salt, common name mecoprop. This compound causedsevere epinasty and stimulated the production of ethylene fromshoot explants. However, when intact plants were treated withethylene, the leaves became only slightly epinastic. The ethyleneprecursor, 1-aminocyclopropane-I-carboxylic acid (ACC), at concentrationswhich stimulated the release of ethylene, was equally ineffectivein causing epinasty. Furthermore, 2, 5-norbornadiene, a specific,competitive inhibitor of ethylene action, only partly alleviatedmecoprop-induced epinasty. The responses observed in chickweedwere compared with those produced in tomato plants. ACC inducedepinasty in tomato within 2 h and these symptoms were completelyinhibited by norbornadiene. However, as in chickweed, the inhibitorgave only partial reversal of mecoprop-induced epinasty, implyingthat the epinastic response caused by the herbicide was notattributable to ethylene alone. We therefore suggest that mecoprop-inducedepinasty is a result of the combined ethylene-stimulating andgrowth-promoting properties of the herbicide. Mecoprop-stimulated ethylene evolution was initially significantlygreater in a herbicide-resistant, compared with a more susceptiblebiotype of chickweed. The significance of this finding is discussedin relation to the mechanism of mecoprop resistance in chickweed. Epinasty, ethylene, (RS)-2-(4-chloro-o-tolyloxy)propionic acid, mecoprop, herbicide resistance, chickweed, Stellaria media L., tomato, Lycopersicon esculentum L.     

The Mode of Flower-Inhibiting Action of Ethylene in Pharbitis nil

The inhibitory effect of ethylene on photoperiodic flower inductionin Pharbitis nil was investigated in relation to the time ofethylene application. Ethylene applied during an 18-h dark periodnot only made the dark period non-inductive, but also greatlyinhibited the flower-inducing effect of the 2nd 18-h dark periodgiven 6 h after the end of the first dark period. The seconddark period was inductive when it was given 30 h after the endof the first dark period, during which ethylene was applied.Ethylene applied during the light period prior to an inductivedark period had no inhibitory effect, suggesting that ethylenegiven during the dark period produces some flower-inhibitingentity. (Received April 17, 1987; Accepted June 17, 1987)     

Shoot Auxin and Light in the Mechanism Maintaining the Posture of the Cotyledons of Linum usitatissimum

The mean cotyledon angle (liminal day posture) of flax is linearlyrelated to light intensity, the cotyledons being raised withincreasing intensity; adjustments occur when plants are transferredto new conditions. Short days, low light intensity, and prolongeddarkness result in epinasty. Epinastic and hyponastic responsesare enhanced by excision of the epicotyl shoot. Applicationof indoleacetic acid to the cut shoot is an effective substitutefor the excised portion. Ringing the intact epicotyl with 2,3,5-tri-iodobenzoicacid has an effect comparable to excision. Given continuously or in a light/dark cycle, blue light induceshyponasty, and red light maintains the liminal posture for about48 h after which time epinasty occurs. Far-red light given continuouslyinduces epinasty to an extent comparable to the effect of darkness.Irradiation with far-red light even for brief periods priorto placing the plants in darkness enhances the epinastic response:the effect of far-red light is reversed by a subsequent shortexposure to red light. Interruption of a prolonged period of darkness by blue lightconsiderably lessens the subsequent epinastic reaction whetheror not any epinasty has occurred before the light-break. Redor white light given under these circumstances has less effect.     

An Evaluation of the Effects of Ethylene on Carbon Assimilation in Lycopersicon esculentum Mill

Leaf and whole plant gas exchange rates of Lycopersicon esculentumMill, were studied during several days of continuous exposureto ethylene. Steady-state photosynthesis and transpiration ratesof control and ethylene-treated individual leaves were equivalent.However, the photosynthesis and transpiration rates of treatedleaves required at least five times longer to reach 50% of thesteady-state rate. This induction lag was attributed to ethylene—inducedleaf epinasty and temporary acclimation to lower incident lightlevels immediately prior to measurement of gas exchange. Thewhole plant net carbon exchange rate (NCER) of a representativetreated plant was also reduced by 51% after 24 h exposure toethylene relative to both its pre-treatment rate and that ofthe control. Ethylene exposure reduced the growth rate of thetreated plant by 50% when expressed as carbon (C) gain. Theinhibition of NCER and growth rate associated with epinastywas completely reversed when the epinastic leaves were returnedto their original positions and light interception was re-established.The results demonstrate that the inhibition of whole plant CO₂assimilation is indirect and due to reduced light interceptionby epinastic leaves. Morphological changes caused by environmentalethylene are thus shown to reduce plant C accumulation withoutinhibiting leaf photosynthesis processes per se. Key words: Ethylene, carbon assimilation, growth     

Ethylene, seed germination, and epinasty

Ethylene activity in lettuce seed (Lactuca satina) germination and tomato (Lycopersicon esculentum) petiole epinasty has been characterized by using heat to inhibit ethylene synthesis. This procedure enabled a separation of the production of ethylene from the effect of ethylene. Ethylene was required in tomato petioles to produce the epinastic response and auxin was found to be active in producing epinasty through a stimulation of ethylene synthesis with the resulting ethylene being responsible for the epinasty. In the same manner, it was shown that gibberellic acid stimulated ethylene synthesis in lettuce seeds. The ethylene produced then in turn stimulated the seeds to germinate. It was hypothesized that ethylene was the intermediate which caused epinasty or seed germination. Auxin and gibberellin primarily induced their response by stimulating ethylene production.     

Ethylene accumulation in waterlogged Rumex plants promotes formation of adventitious roots

Accumulation of the gaseous plant hormone ethylene is very importantfor the induction of several responses of plants to flooding.However, little is known about the role of this gas in the formationof flooding-induced adventitious roots. Formation of adventitiousroots in Rumex species is an adaptation of these plants to floodedsoil conditions. The large air-spaces in these roots enablesdiffusion of gases between shoot and roots. Application of ethylene to non-flooded Rumex plants resultedin the formation of adventitious roots. In R. palustris Sm.shoot elongation and epinasty were also observed. The numberof roots in R. thyrsiflorus Fingerh. was much lower than inR. palustris, which corresponds with the inherent differencein root forming capacity between these two species. Ethyleneconcentrations of 1.5–2µI I_{– 1} induced a maximumnumber of roots in both species. Quantification of ethylene escaping from root systems of Rumexplants that were de-submerged after a 24 h submergence periodshowed that average ethylene concentrations in submerged rootsreached 1.8 and 9.1 µl I_–1 in R. palustris and R.thyrsiflorus, respectively. Inhibition of ethylene productionin R. palustris by L--(2-aminoethoxyvinyl)-glycine (AVG) or-aminobutyric acid (AIB) decreased the number of adventitiousroots induced by flooding, indicating that high ethylene concentrationsmay be a prerequisite for the flooding-induced formation ofadventitious roots in Rumex species. Key words: Adventitious roots, epinasty, ethylene, flooding, Rumex, shoot elongation     

Brassinosteroid-induced epinasty in tomato plants

The effects of root treatments of brassinosteroid (BR) on the growth and development of hydroponically grown tomato plants (Lycopersicon esculentum Mill cv Heinz 1350) were evaluated. There was a dramatic increase in petiole bending when the plants were treated with 0.5 to 1.0 micromolar BR. The leaf angle of the treated plants was almost three times that of untreated controls. BR-induced epinasty appeared to be due to stimulation of ethylene production. Excised petioles from BR-treated plants produced more than twice as much ethylene as did untreated controls. As ethylene production increased, the degree of petiole bending also increased, and inhibition of ethylene production by AOA or CoCl₂ also inhibited epinasty. BR-treated plants had increased levels of the ethylene precursor 1-aminocyclopropane-1-carboxylic acid (ACC) in the leaf tissue. ACC appeared to accumulate primarily in the petioles with the greatest amount of ACC accumulating in the youngest petioles. Time course evaluations revealed that BR treatment stimulated ACC production. As ACC accumulated, ethylene increased, resulting in epinasty. Little or no ACC was found in the xylem sap, indicating that there was a signal transported from the roots which stimulated ACC synthesis in the leaf tissue.     

Auxin and Ethylene Regulation of Petiole Epinasty in Two Developmental Mutants of Tomato, diageotropica and Epinastic

The epinastic growth responses of petioles to auxin and ethylene were quantified in two developmental mutants of tomato (Lycopersicon esculentum Mill.). In the wild type parent line, cultivar VFN8, the epinastic response of excised petiole sections was approximately log-linear between 0.1 and 100 micromolar indole-3-acetic acid (IAA) and 2,4-dichlorophenoxyacetic acid (2,4-D) concentrations, with a greater response to 2,4-D at any concentration. When ethylene synthesis was inhibited by aminoethoxyvinylglycine (AVG), epinasty was no longer induced by auxin, but could be restored by the addition of ethylene gas. In the auxin-insensitive mutant, diageotropica (dgt), no epinastic response to IAA was observed at IAA concentrations that effectively induced epinasty in VFN8. In the absence of added IAA, epinastic growth of dgt petioles in 1.3 microliters per liter exogenous ethylene gas was more than double that of VFN8 petioles. IAA had little additional effect in dgt, but promoted epinasty in VFN8. These results confirm that tomato petiole cells respond directly to ethylene and make it unlikely that the differential growth responsible for epinasty results from lateral auxin redistribution. The second mutant, Epinastic (Epi), exhibits constitutively epinasty, cortical swelling, and root branching symptomatic of possible alternation in auxin or ethylene regulation of growth. Only minor quantitative differences were observed between the epinastic responses to auxin and ethylene of VFN8 and Epi. However, in contrast to VFN8, when ethylene synthesis or action was inhibited in Epi, auxin still induced 40 to 50% of the epinastic response observed in the absence of inhibitors. This indicates that the target cells for epinastic growth in Epi are qualitatively different from those of VFN8, having gained the ability to grow differentially in response to auxin alone. The dgt and Epi mutants provide useful systems in which to study the genetic determination of target cell specificity for hormone action.     

Effects of root anaerobiosis on ethylene production, epinasty, and growth of tomato plants

Experiments were performed to determine the source(s) of ethylene-causing epinasty in flooded tomato plants (Lycopersicon esculentum Mill.). Simultaneous measurements were made of ethylene synthesized by the roots and shoots of tomato plants exposed to either aerobic or anaerobic atmospheres in the root zone. When the root zone was made anaerobic by a flowing stream of N₂ gas, petiole epinasty and accelerated ethylene synthesis by the shoots were observed. In soil-grown plants, ethylene synthesis by the root-soil complex increased under anaerobic conditions; but when grown in inert media under the same conditions, ethylene synthesis by roots remained constant or declined during the period of rapid epinastic growth by the petioles. Other characteristic symptoms of flooding, e.g. reduced growth and chlorosis, were also observed in plants with anaerobic roots. Pretreatment of plants with AgNO₃, an inhibitor of ethylene action, completely prevented epinasty, demonstrating that ethylene is the agent responsible for waterlogging symptoms. These results indicate that deprivation of O₂ to the roots is the primary effect of soil flooding, and that this is sufficient to cause increased ethylene synthesis in the shoot. The basis of the observed root-shoot communication is unknown, but root-synthesized hormones or specific ethylene-promoting factors may be involved.     

Failure of Ethylene to Change the Distribution of Indoleacetic Acid in the Petiole of Coleus blumei X frederici during Epinasty

The effect of ethylene on the distribution of applied indoleacetic acid in the petiole of Coleus blumei Benth. X C. frederici G. Taylor has been investigated during the development of epinastic curvature. Using intact plants, ¹⁴C-IAA was applied to the distal region of the leaf lamina and the accumulation of label in the abaxial and adaxial halves of 5 mm petiole sections was determined after 1.5, 3, and 6 hours. Over this period the label was transported out of the lamina into the petiole at a rate of at least 66 mm hr⁻¹. Of the total amount of label in the petiole sections, 24 to 30% was located in the adaxial half and this distribution was not altered significantly by exposing plants to an atmosphere containing 50 μl/l ethylene. Thus when epinastic curvature is induced by ethylene there is no associated increase in the IAA content of the expanding adaxial half. The role of endogenous IAA in petiole epinasty was studied by restricting its movement with DPX 1840 (3,3a-dihydro-2-[p-methoxyphenyl]-8H-pyrozolo{5,1-a}isoindol-8-one). The leaf petioles still showed an initial epinastic response to ethylene. It is concluded that ethylene-induced epinasty is not dependent upon either any change in the transport of IAA or its redistribution within the petiole.     

Ethylene-induced opposite redistributions of calcium and auxin are essential components in the development of tomato petiolar epinastic curvature.

Calcium has been suggested as an important mediator of gravity signaling transduction within the root cap statocyte. In a horizontally-placed root, it is redistributed in the direction of the gravity vector (i.e. it moves downward) and its redistribution is closely correlated with auxin downward movement. However, the involvement of calcium in the regulation of ethylene-induced epinasty and auxin movement is not known. In this report, we examined the involvement of calcium in lateral auxin transport during ethylene-induced epinasty in an effort to understand the relationship among calcium, auxin, and ethylene. Ethylene-induced epinasty was further stimulated by exogenously applied Ca2+, the calcium effect being the strongest among divalent cations tested. Pretreatment with NPA, an auxin transport inhibitor, negated the promotive effect of calcium ions on the petiolar epinasty. Ethylene caused redistribution/differential accumulation of 45Ca2+ toward the morphologically lower (abaxial) side of the leaf petioles, an effect opposite to that of 14C-IAA redistribution. Verapamil, a Ca2+ channel blocker, inhibited ethylene-induced epinasty, as well as the redistribution of 14C-IAA and 45Ca2+. When the petiole was inverted in the presence or absence of ethylene, the direction of 45Ca2+ differential accumulation was still toward the morphologically abaxial side of the petiole during epinastic movement regardless of gravitational direction. These results suggest that gravity-insensitive, ethylene-induced Ca2+ redistribution and accumulation toward the abaxial side are closely coupled to the adaxial auxin redistribution/accumulation and, in turn, to the petiolar epinasty.     

Relation between Sex Expression Types and Cotyledon Etiolation of Cucumber in vitro I. On the Role of Ethylene Evolved from Seedlings

Cucumber seedlings, when cultured in vitro, showed differencesin cotyledon etiolation rates among cultivars with differentgenetic backgrounds for sex expression. The chlorophyll contentin gynoecious cultivars (acr^F/acr^F) decreased rapidly whilethat in monoecious ones (acr⁺/acr⁺) decreased more slowly, andthat in mono-gynoecious ones (acr¹/acr¹) decreased at an intermediaterate. Etiolation was suppressed even in early-etiolating cultivarswhen the flask remained unsealed or endogenously evolved ethylenewas removed. Cotyledon etiolation was enhanced even in late-etiolatingcultivars when ethephon was added to the flask. The rate ofetiolation corresponded to the ethylene concentration in theflask; much more ethylene was detected in early-etiolating cultivarsthan in late-etiolating ones. Ethylene accumulation is one of the important factors involvedin the cotyledon etiolation observed in in vitro cultures. Thedifference in etiolation rates among seedlings with differentgenetic backgrounds for sex expression corresponds to theirability for ethylene evolution, in the order of acr^F>acr¹>acr⁺. (Received January 6, 1981; Accepted March 23, 1981)     

Auxin-Induced Epinasty of Tobacco Leaf Tissues (A Nonethylene-Mediated Response)

Interveinal strips (10 x 1.5 mm) excised from growing tobacco (Nicotiana tabacum L. cv Xanthi) leaves curled >300[deg] when incubated for 20 h in 5 to 500 [mu]M [alpha]-naphthalene acetic acid or 50 to 500 [mu]M indole-3-acetic acid. Epinasty was not induced without auxin or by the auxin analog [beta]-naphthalene acetic acid, and less substantial epinasty was induced in midrib and vein segments. Auxin treatment increased the length of both surfaces of strips. Curvature resulted from greater growth on the adaxial side. Epinastic sensitivity of strips to auxin appeared first in the distal third of young leaves (blade 4.5-6.0 cm). In older leaves (8-10 and 12-14 cm), the interveinal tissues throughout were sensitive, whereas in leaves 16- to 18-cm long, sensitivity was reduced in the distal two-thirds. Amino-oxyacetic acid (AOA), an ethylene biosynthesis inhibitor, partially inhibited epinasty at 100 [mu]M. However, a poor correlation between inhibition of ethylene biosynthesis by AOA and its inhibition of curvature and the inability of ethylene to produce epinasty or to reverse the effects of AOA suggests that auxin-induced epinasty is not caused by auxin-induced ethylene production.     

Prevention of salt-induced epinasty by α-aminooxyacetic acid and cobalt

Ethylene production in leaf petiole and laminae tissues was stimulated in tomato (Lycopersicon esculentum Mill. cv. UCT5) plants exposed to salinity-stress. At the highest salinity level (250 mM NaCl), rates of ethylene production more than doubled over those observed in non-stressed plants. Correspondingly, petiolar epinasty increased with increasing levels of stress impositions. Both responses were suppressed when either 1 mM -aminooxyacetic acid (AOA), or 100 M Co²⁺ was simultaneously applied. Co²⁺, but not AOA, had a pronounced effect on ethylene production resulting from the application of a saturating dose (2 mM) of 1-aminocyclopropane-1-carboxylic acid (ACC), the immediate precursor of ethylene. This result suggests that ethylene production is dependent upon the activity of ethylene forming enzyme (EFE). The magnitude of ethylene stimulation in leaf petioles was related to the salinity level imposed and to the induction of petiole epinasty. In the absence of stress impositions, epinastic responsiveness to ethylene or its precursor, ACC, might provide a simple, indirect criteria to adjudge salt-sensitivity among plants.Research supported by AID contract II, NEB-1070-A-00-2074-00.     

Obtention d'épinastie des cotylédons de Sinapis alba en éclairement rouge Iointain continu.

Obtention of epinasty in the cotyledons of Sinapis alba under continuous far-red illumination. Relation to the development of hypocotyl and cotyledons. Treatment by far red light induces epinasty of the cotyledonary leaf. The effects of light treatments of different intensities were recorded for hypocotyl length, percentage of epinasty and the cotyledon leaf area. The percentage of epinasty increases as a function of treatment duration and light intensity. As from the second day of treatment a correlation is obtained between the percentage of epinasty and hypocotyl length when the seedlings are given far-red light treatment of different durations.     

Interpreting Plant Responses to Clinostating: I. MECHANICAL STRESSES AND ETHYLENE

The severe epinasty and other symptoms developed by clinostated leafy plants could be responses to gravity compensation and/or the mechanical stresses of leaf flopping. Epinasty in cocklebur (Xanthium strumarium L.), tomato (Lycopersicon esculentum Mill.), and castor bean (Ricinus communis L.) is delayed by inhibitors of ethylene synthesis and action (aminoethoxyvinylglycine and Ag⁺), confirming the role of ethylene in clinostat epinasty. To test the possibility that clinostat mechanical stresses (leaf flopping) cause ethylene production and, thus, epinasty, vertical plants were stressed with constant, gentle, horizontal, or vertical shaking or with a quick, back-and-forth rotation (twisting). Clinostat leaf flopping was closely approximated but with a minimum of gravity compensation, by turning plants so their stems were horizontal, rotating them quickly about the stem axis, and then returning them to the vertical, repeating the treatment every four minutes (clinostat rotation time). None of these mechanical stresses produced significant epinasties, but vigorous hand-shaking (120 seconds per day) generated minor epinasties, as did Ag⁺ applied daily (concentrations high enough to cause leaf browning). Plants gently inverted every 20 minutes developed epinasty at about the same rate and to about the same extent as clinostated plants, but plants inverted every 20 minutes and immediately returned to the upright position did not become epinastic. It is concluded that clinostat epinasty is probably caused by disturbances in the gravity perception mechanism, rather than by leaf flopping.     

油莱素甾醇促进黄化水稻第二叶切段偏上性生长与乙烯的关系

油菜素甾醇强烈刺激黄化水稻第二叶切段倾斜的实质是偏上性生长。油菜素甾醇处理的切段的叶枕细胞带发生了不均一的可塑性生长,叶枕薄壁细胞明显增大。油菜素甾醇促进切段偏上性生长的同时伴随着乙烯释放。两促进效应能被乙烯生物合成抑制剂CoCL2平行抑制,表明两效应存在正相关。     

Auxin and Ethylene-stimulated Adventitious Rooting in Relation to Tissue 9

We have previously shown that both endogenous auxin and ethylenepromote adventitious root formation in the hypocotyls of derootedsunflower (Helianthus annuus) seedlings. Experiments here showedthat promotive effects on rooting of the ethylene precursor,1-aminocyclopropane-l-carboxylic acid (ACC) and the ethylene-releasingcompound, ethephon (2-chloro-ethylphosphonic acid), dependedon the existence of cotyledons and apical bud (major sourcesof auxin) or the presence of exogenously applied indole-3-aceticacid (IAA). Ethephon, ACC, aminoethoxyvinylglycine (an inhibitorof ethylene biosynthesis), and silver thiosulphate (STS, aninhibitor of ethylene action), applied for a length of timethat significantly influenced adventitious rooting, showed noinhibitory effect on the basipetal transport of [³H]IAA. Theseregulators also had no effect on the metabolism of [³H]IAA andendogenous IAA levels measured by gas chromatography-mass spectrometry.ACC enhanced the rooting response of hypocotyls to exogenousIAA and decreased the inhibition of rooting by IAA transportinhibitor, N-1-naphthylphthalamic acid (NPA). STS reduced therooting response of hypocotyls to exogenous IAA and increasedthe inhibition of rooting by NPA. Exogenous auxins promotedethylene production in the rooting zone of the hypocotyls. Decapitationof the cuttings or application of NPA to the hypocotyl belowthe cotyledons did not alter ethylene production in the rootingzone, but greatly reduced the number of root primordia. We concludethat auxin is a primary controller of adventitious root formationin sunflower hypocotyls, while the effect of ethylene is mediatedby auxin. Key words: Auxin, ethylene, adventitious rooting, sunflower     

Evidence that Ethylene is not Involved in Red-light-induced Stimulation of Chlorophyll Formation in Etiolated Cucumber Cotyledons

The involvement of ethylene in red-light-induced stimulationof chlorophyll (Chl) formation was studied because one of thered-light effects on Chl formation (the lateappearing effect)interacts with the ethylene effect in 3-day-old excised etiolatedcotyledons of cucumber (Cucumis sativus L. cv. Aonagajibai).Ethylene production by etiolated cotyledons of intact seedlingsin the dark is enhanced by a red-light pulse, but the effectdoes not occur in excised cotyledons. Application of ethylenein the dark to 3-day-old intact seedlings has little effecton Chl formation in the cotyledon during subsequent continuousillumination, although ethylene pretreatment of 5-day-old seedlingssignificantly stimulates Chl formation. Removal of endogenousethylene by mercuric perchlorate [Hg(ClO₄)₂] does not specificallysuppress the red-light action on Chl formation in both attachedand excised cotyledons. Inhibition of ethylene synthesis byaminoethoxyvinylglycine does not affect the red-light effecton Chl formation in excised cotyledons. These facts indicatethat ethylene does not operate as a mediator of red light instimulating Chi formation in either attached or excised cotyledons. (Received December 13, 1981; Accepted March 30, 1981)     

Transport of shoot- and cotyledon-applied indole-3-acetic acid to Vicia faba root

To clarify the participation of indole-3-acetic acid (IAA) originatingfrom the shoot in root growth regulation and the mechanism ofIAA translocation from shoot to root, the movement of ¹⁴C-IAAwhich was applied to the epicotyl or the cotyledon of Viciafaba seedlings was investigated. The radioactivity of IAA appliedto the cotyledon moved faster to the root tip than that appliedto the epicotyl. On the basis of the effect of 2,3,5-triiodobenzoic acid on IAAmovement, a comparison with ¹⁴C-glucose movement and autoradiographicexamination, the nature of IAA movement was concluded to bepolar transport from the epicotyl to the basal part of the roots,while IAA movement from the epicotyl to the cotyledon, fromthe basal part of roots to the apical part, and from the cotyledonto the epicotyl and to the root took place in the phloem. Theradioactivity from ¹⁴C-IAA applied to the cotyledon accumulatedin lateral root primordia and vascular bundles. These factssuggest that IAA produced in cotyledons may participate in theregulation of Vicia root development. (Received December 21, 1979; )