A comparative study of the response of the roots and shoots of sunflower and maize to mechanical stimulation

Despite numerous studies of the effects of mechanical stimulationon plant shoots, the response of roots to mechanical stimulationhas largely been neglected. In this study the effects of shootflexure on the morphology and mechanics of two contrasting speciesof herbaceous angiosperm, growing in a glasshouse were compared:maize (Zea mays), a monocot; and sunflower (Helianthus annuusL.) a dicot. Mechanical stimulation affected the root more than the shootcomponents. Root systems of mechanicallystressed sunflowershad a greater angle of spread and increased root number. Aswell as large morphological and weight effects, with increasesover the control of 33% in the length of rigid root and 38%in the dry weight of lateral roots, in sunflowers, there werealso mechanical effects. In both species roots of flexed plantswere more rigid, stronger and composed of stiffer material andtheir root systems also provided greater anchorage strength.In contrast, there was only a small reduction in shoot weightand shoot height in flexed plants and no effects on mechanicalproperties. There were differences in behaviour between species; maize rootmorphology responded less than that of sunflowers to mechanicalstimulation. The basal diameter of roots increased by only 8%compared with 16% in sunflowers, though the roots of both speciesshowed similar increases in material stiffness. This differenceis related to the lack of secondary thickening in the monocotscompared with the dicot sunflowers. Key words: Thigmomorphogenesis, Helianthus annuus L., Zea mays, anchorage, lodging     

The Responses of Field-grown Sunflower and Maize to Mechanical Support

The effects of mechanical support on two contrasting speciesof herbaceous annual, the dicot sunflower (Helianthus annuusL.) and the monocot maize (Zea mays L.), were investigated bycomparing the growth and mechanical properties of supportedplants and those which were left to sway freely in the wind. Providing support had its greatest effect on the more highly-stressedbasal areas of the plants, such as the lower stem and the baseof the lateral roots. The diameter of the stem bases of bothspecies was approx. 10% lower in supported plants, but therewas no difference between treatments in the diameter of thestem above 50 cm. Roots of both species also showed a reductionin rigidity and bending strength of 40–50% in the supportedplants compared with freely swaying plants. There was a significantreduction in the partitioning of biomass to the root systemsof supported plants of both species. There were differences in the way in which sunflower and maizeresponded to the provision of support; in sunflower, the reductionin lateral diameter was about twice that in maize, whereas inmaize the decrease in the number of first-order laterals wastwice that of sunflower. This study suggests that thigmomorphogenesismay be a localized response, but that different species canrespond in different ways to mechanical stimulation. Wind; support; anchorage; thigmomorphogenesis; Helianthus annuus L.; sunflower; Zea mays L.; maize     

The Effects of Soil Bulk Density on the Morphology and Anchorage Mechanics of the Root Systems of Sunflower and Maize

The effects of soil bulk density and hence strength on two contrastingspecies of herbaceous annuals, the dicot sunflower (HelianthusannuusL.) and the monocot maize (Zea maysL.), were investigatedby comparing the morphology and mechanics of field-grown plantsin soil with a low and high bulk density. Soil with a low bulkdensity had a significantly lower penetration resistance (118±4.4kPa) than the high bulk density soil (325±12.2 kPa;P<0.0001).Soil strength affected shoot and root systems of both speciesbut had no significant effect on shoot height. In both speciesroots were thicker closer to the stem base in strong soil comparedto those in weaker soil. Sunflower tap-roots growing in strongsoil tapered more rapidly than those in weak soil. Only in maize,however, were roots growing in weak soil stiffer than thosein strong soil. Despite only small absolute differences in thepenetration resistance of the soil both species growing in strongsoil had greater anchorage strength than those in weak soil.As a consequence more plants in weak soil lodged compared withthose growing in strong soil. This study shows that plants can,to a small extent, respond to changes in soil strength, butthat changes do not appear to compensate fully for alterationsin soil conditions. Furthermore it may be possible, by manipulatingsoil strength, to control lodging.Copyright 1999 Annals of BotanyCompany Roots, compaction, soil strength, anchorage mechanics, bulk density, thigmomorphogenesis, lodging,Helianthus annuusL.,Zea maysL.     

Profiles of Leaf Senescence During Reproductive Growth of Sunflower and Maize

We investigated the effect of reproductive growth on the profilesof leaf senescence in maize (Zea mays L.) and sunflower (Helianthusannuus L.). Leaf senescence after flowering was assessed usingboth structural (leaf chlorophyll, nitrogen and dry matter)and functional (photosynthesis) variables in undisturbed plants(+G) and in plants in which grain set was prevented (-G). Twoweeks after flowering, lack of grain accelerated senescencein maize and delayed senescence in sunflower as indicated byleaf chlorophyll; leaf nitrogen and dry matter were less sensitiveresponse variables. Lack of interaction between reproductivetreatment and leaf position indicates that the senescence signal,whatever its nature, was equally effective throughout the plantin both species. In both species, feedback inhibition of photosynthesiswas first detected 30–35 d after flowering; excess carbohydratein the leaves was therefore an unlikely trigger of acceleratedsenescence in maize. As reproductive development progressed,differences between +G and -G plants were more marked in sunflower,and tended to disappear or reverse in maize. In sunflower, interactionsbetween leaf position and reproductive treatment—attributableto the local effect of grain—were detected around 20–27d after flowering. Copyright 2000 Annals of Botany Company Helianthus annuus, Zea mays, chlorophyll, light, nitrogen, photosynthesis, reproductive growth, senescence, source-sink, SPAD.     

Male Sterility and Anther Wall Structure in Copper-deficient Plants

Anther development and pollen sterility were followed in plantsof wheat, oat, barley, sweetcorn, sunflower, petunia and subterraneumclover grown at a range of copper supplies. Copper-deficientplants had increased pollen sterility. Lignified wall thickenings were reduced or absent in the endotheciaof anthers from Cu-deficient plants. Reduced seed set may resultboth from reduced pollen fertility or failure of the stomiato rupture due to decreased lignification of anther walls. Triticum aestivum L., wheat, Hordeum vulgare L., barley, Avena sativa L., oat, Zea mays L., corn, sweetcorn, maize, Helianthus annuus L., sunflower, Petunia hybrida L., Trifolium subterraneum L., subterranean clover, male sterility, anther development, copper deficiency     

A Comparative Study of the Anchorage Systems of Himalayan Balsam Impatiens glandulifera and Mature Sunflower Helianthus annuus

The anchorage systems of Himalayan balsam Impatiens glanduliferaand mature sunflowers Helianthus annuus were investigated bycombining morphological and anatomical study of the root systemswith mechanical tests on roots and with studies in which matureplants were pulled over. The root system of balsam is dominated by large numbers of fleshytapering adventitious roots which point downwards from theirorigin at the wide stem base. Sunflowers, in contrast, havea tapering tap-root from which 20–30 well-branched lateralsemerge, pointing radially outwards and downwards. Roots of eachspecies have contrasting anatomy: those of balsam resemble stems,having a central watery pith and being strengthened peripherallyby lignification of vascular tissue; roots of sunflowers arestrengthened by a solid woody stele. Roots of both species arerigid in tension and, towards the base, in bending. Both species exhibited similar behaviour to that known for treessuch as Sitka spruce; when pulled over they rotated about ahinge leeward of the stem base and a root-soil ball was pulledout of the surrounding soil. Anchorage was resolved into threecomponents which, in order of decreasing magnitude, were (i)the resistance to pulling of the roots on the windward sideof the plant (and, for sunflower, the tap-root); (ii) the resistanceof roots and soil at the leeward hinge to rotation; and (iii)the weight of the root-soil ball. Sunflower had stronger anchoragebut achieved it at a greater cost in terms of the dry mass ofits root system. In each species, the morphology, anatomy and mechanical propertiesof the root system can be related to those of the stem. Thewide stem base of balsam allows large numbers of mechanicallyefficient fleshy roots to be attached whereas in sunflowersa woody tap-root system is necessary to anchor the much narrowerstem. Key words: Impatiens, Helianthus, roots, anchorage     

Does ABA in the Xylem Control the Rate of Leaf Growth in Soil-Dried Maize and Sunflower Plants?

Maize (Zea mays L.) and sunflower (Helianthus annuus L.) plantswere grown in large volumes of soil and leaf growth rate wasmonitored on a daily basis. Half the plants were given a soildrying treatment and when they showed a significant restrictionof growth rate (compared to both their daily growth rate beforedrying and the average growth rate of well-watered plants onthe same day), leaf water relations were measured and xylemsap was extracted using several techniques. There was a significant negative log-linear relationship betweenthe rate of leaf growth and the concentration of ABA in thexylem for both species. There was no clear relationship betweenleaf growth rate and leaf water potential or turgor for eitherspecies. Assessment of different methods for sampling xylemsap suggests that exudates collected from stem stumps or samplescollected by pressurizing the whole root system are suitablefor estimating ABA concentration in xylem, at least with largeplants of maize or sunflower, provided the first few hundredcubic millimetres of collected sap are used for the assay. Centrifugationof sections of stems resulted in dilution of ABA in the xylemsap with sap squeezed from parenchyma tissue. This is because,at least in plants subjected to mild soil drying, the concentrationof the ABA in the xylem is far higher than that in the cellsap of stem tissue. Results support the proposal that ABA plays a major role asa chemical signal involved in the root-to-shoot communicationof the effects of soil drying. The non-hydraulic restrictionof leaf growth by a chemical signal can be explained by theextra root-sourced ABA in the xylem and may be an importantcomponent of the modification of growth and development whichresults from prolonged soil drought. Key words: Soil drying, ABA, leaf growth, Zea mays L., Helianthus annuus L.     

Adaptation and Functional Balance Between Shoot and Root Activity of Sunflower Plants Grown at Different Root Temperatures

Helianthus annuus L. plants were grown with the shoots at normalair temperature and with the roots in nutrient solution at 10,20 or 30 °C. The higher the root temperature the greaterthe growth of the leaves, resulting in higher production ofphotosynthates. Irrespective of growth conditions an equilibriumwas established between the maintenance respiratory activityof shoot and roots. A constant proportion of the photosynthateproduced was used in respiration. The results are discussedin relation to a thermodynamic theory of stability of biologicalsystems. It is suggested that changes in energy partition betweenmaintenance and growth, and then in relative growth rates ofshoots and roots during an adaptation period, represent a majorhomeostatic mechanism. shoots, roots, maintenance respiration, growth, relative growth rate, respiration, adaptation, sunflower, Helianthus annuus L.     

Relationship Between Root Morphogenesis and Period of Predominant Cell Arrest in Intact and Aseptically-cultured Roots of Helianthus annuus L.

Root morphogenesis and cell cycle kinetics of intact and aseptically-grownexcised roots of Helianthus annuus L. were studied. Intact rootsshow predominant cell arrest in G1 with an absence of polyploidcells coincident with secondary vascularization. Exposure ofthe cut ends of aseptically grown excised roots to known concentrationsof indol-3-yl acetic acid, benzyladenine, and myo-inositol for8 weeks initiated the production of secondary vascular tissuesand predominant cell arrest in G2 concommitant with poiyploidization.Excised roots grown in the absence of these substances producedroots with only primary vascularization and predominant cellarrest in G1 coincident with an absence of polyploidization.These results indicate that (a) root cells of H. annuus havethe ability to undergo polyploidization that may be inducedby exogeneously applied chemicals, (b) a general relationshipbetween predominant cell arrest in G1 coincident with the absenceof secondary vascularization does not hold true and (c) althoughsecondary vascularization occurs in cultured roots exposed toall three additives similar to secondary vascularization inintact roots, the two roots should not be considered identicalin all respects. Helianthus annus L., sunflower, root, morphogenesis, cell cycle kinetics, polyploidy, cell differentiation, vascularization     

Distribution of Protein-bound Hexosamine in Chloroplasts

Intact chloroplasts of spinach (Spinacia oleracea L.), sunflower (Helianthus annuus L.), and maize (Zea mays L.) mesophyll cells contained 0.33, 0.50, and 0.14% of bound hexosamine on a protein basis, respectively. Undifferentiated maize chloroplasts contained 0.19%. Values for chloroplast lamellae were, respectively, 0.16, 0.18, 0.12, and 0.06% and for envelope membranes they were 1.6, 2.5, 3.8, and 2.7%. Thus most of the hexosamine of chloroplasts is located in the envelope membrane.     

The uptake of persistent organic pollutants by plants

In a field experiment, the transfer of polychlorinated biphenyls (PCBs) and polycyclic aromatic hydrocarbons (PAHs) from contaminated soil to maize (Zea mays L.), sunflower (Helianthus annuus), poplar (Populus nigra × P. maximowiczii) and willow (Salix × smithiana) and the distribution of PCB congeners in maize and sunflower was investigated. The former waste incinerator in Hradec Králové (Czech Republic) was chosen for the experiment. Results of plot screening showed heterogenous contamination by PCBs and PAHs. PCB soil contamination was evidently caused by Delor 106 or Aroclor 1260 stocking and PAH contamination by chemicals containing fluoranthene, benzo/b/fluoranthene, phenanthrene and pyrene. Tested plants were planted on a contaminated field site, in soil contaminated with 1530 μg/kg of total PCBs and 0.138 and 3.42 mg/kg of total PAHs. The results show that maize and sunflower roots accumulated the most PCBs from soil. These plants accumulated hexa- and heptachlorobiphenyl congeners more than tri-, tetra-, and pentachlorobiphenyl congeners. Total concentrations of PAHs in tested plants ranged from 0.096 to 1.34 mg/kg. The highest phenanthrene concentration was found in aboveground biomass of sunflower and the highest concentration of pyrene, in maize roots.     

Effects of Water Stress and High-Temperature Stress on the Structure and Activity of Photosynthetic Apparatus of Zea Mays and Helianthus Annuus

Effects of high-temperature stress (HTS) and PEG-induced water stress (WS), applied separately or in combination, on the functional activity and ultrastructure of the photosynthetic apparatus (PSA) of maize (Zea mays L.) and sunflower (Helianthus annuus L.) plants were investigated. In maize plant tissues WS provoked the decrease in RWC by 10.9 %, HTS by 7.0 %, and after simultaneous application of the both treatments the decrease was 32.7 % in comparison with control plants. Similar but more expressed changes were observed in sunflower plants. Sunflower was more sensitive to these stresses. Net photosynthetic rate decreased significantly after all treatments, more in sunflower. In mesophyll chloroplasts after separately applied WS and HTS the number of grana and thylakoids was reduced and electron-transparent spaces appeared. At combined stress (WS+HTS) granal and stromal thylakoids were considerably affected and chloroplast envelope in many of them was partially disrupted. This revised version was published online in August 2006 with corrections to the Cover Date.     

The Anchorage Mechanics of Maize, Zea mays

The anchorage system of mature maize Zea mays was investigatedby combining morphological and anatomical study of the rootsystem with mechanical tests on roots and with studies in whichplants were pulled over. The root system is dominated by 20–30adventitious roots which emerge in rings from the stem basepointing radially downwards and outwards, approximately 30°from the vertical. Roots are strengthened near their base bya heavily lignified exodermis which makes them rigid in bending;distally, strength and rigidity both decrease because rootsbecome thinner and less lignified. When plants were pulled over,a maximum anchorage moment of 5–20 Nm was mobilized atangles of 8–10°, larger plants having stronger anchorage.Movement was initially centred on the leeward side of the stem,anchorage being due to the resistance of both windward and leewardroots to axial motion through the soil and to bending. At displacementsover 10°, however, leeward roots buckled under combinedbending and compression and the centre of rotation shifted tothe windward perimeter of the root system; subsequent movementof the cone of roots and soil was resisted only by the bearingstrength of the soil beneath it. The differences between anchorage failure in balsam and sunflowersand that in maize probably results from the lower angular spreadand the weakness in compression of the maize roots which preventsthe leeward side of the root system from bearing large downwardloads. The system behaves more like that of wheat; these resultssuggest that the lodging resistance of both plants may be improvedby increasing the bending strength and angle of spread of theadventitious roots. Key words: Zea mays, roots, anchorage     

Structure, Production and Leaf Area Dynamics: A Comparison of Lodgepole Pine and Sunflower

Populations of lodgepole pine (Pinus contorta var latifoliaEngelm) and sunflower (Heltanthus annuus L) were examined tocompare the influence of stand density on canopy structure andthe association between canopy structure and the productionof stem volume The plastic responses of canopy structure tocompetitive interactions indicated that the structural dynamicsof lodgepole pine and sunflower populations were similar, thoughthe degree of plastic response differed The influence of canopystructure on the production of stem volume, however, was fundamentallydifferent efficiency decreased as the mean crown size of lodgepolepine populations increased, but this decrease did not occurin sunflower populations This difference was attributed to thelarge accumulation of branch biomass required to display foliageeffectively in the canopies of the perennial lodgepole pine,accumulation which does not occur in the annual sunflower Lodgepole pine, Pinus contorta var latifolia Engelm, sunflower, Helianthus annuus L, leaf area, canopy structure, stand production, carbon allocation     

Hormone Signals from Roots to Shoots of Sunflower (Helianthus annuus L.). Moderate Soil Drying Increases Delivery of Abscisic Acid and Depresses Delivery of Cytokinins in Xylem Sap

We quantified abscisic acid and a cytokinin trans zeatin ribosideas potential positive or negative signals in root to shoot communicationin sunflower plants (Helianthus annuus L.) growing in dryingsoil. Delivery rates rather than concentration values were usedbecause the former are less subject to change as a result ofdifferences in sap flow through dilution. ABA concentrationand delivery rate increased under the mild drought stress. Incontrast, t-ZR concentration did not change under mild stressalthough delivery rates decreased significantly. With more severedrought stress, both delivery rates and concentration of t-ZRdecreased considerably while ABA concentration and deliverywere enhanced markedly. Root ABA contents mirrored those ofxylem ABA. Helianthus annuus ; soil drying; root signals; ABA; cytokinins; delivery rate; delivery rate     

Lipid and Fatty Acid composition of chloroplast envelope membranes from species with differing net photosynthesis

Lipid and fatty acid compositions were determined for chloroplast envelope membranes isolated from spinach (Spinacia oleracea L.), sunflower (Helianthus annuus L.), and maize (Zea mays L.) leaves. The lipid composition was similar in sunflower, spinach, and undifferentiated maize chloroplast envelope membranes and different in maize mesophyll chloroplast envelope membranes. The predominant lipid constituents in all envelope membranes were monogalactosyldiglyceride (27 to 46%), digalactosyldiglyceride (18 to 33%), and phosphatidylcholine (7 to 30%). The fatty acid composition was also similar in sunflower and spinach chloroplast envelope membranes in comparison to those from maize. The major acyl fatty acids of the chloroplast envelope membrane were palmitic (C_16:0, 41 and 36%) and linolenic (C_18:3, 29 and 40%) acids for spinach and sunflower; palmitic (77%) and stearic (C_18:0, 12%) acids for young maize; and palmitic (61%), stearic (14%), and linolenic (13%) acids for mature maize. The differences in lipid and acyl fatty acid compositions among these plants which vary in their rates of net photosynthesis were largely quantitative rather than qualitative.     

Correlations between proton-efflux patterns and growth patterns during geotropism and phototropism in maize and sunflower

By placing seedlings of sunflower (Helianthus annuus L.) or maize (Zea mays L.) on agar plates containing a pH indicator dye it is possible to observe surface pH patterns along the growing seedling by observing color changes of the indicator dye. Using this method we find that in geotropically stimulated sunflower hypocotyls or maize coleoptiles there is enhanced proton efflux on the lower surface of the organ prior to the initiation of curvature. As curvature develops the pattern of differential acid efflux becomes more intense. A similar phenomenon is observed when these organs are exposed to unilateral illumination, i.e. enhanced acid efflux occurs on the dark side of the organ prior to the initiation of phototropic curvature and the pattern of differential acid efflux intensifies as phototropic curvature develops. These observations indicate that differential acid efflux occurs in response to tropistic stimuli and that the acid efflux pattern may mediate the development of tropistic curvatures.     

Genetic analysis of an electrophoretic variant for the chloroplast-associated form of Cu/Zn superoxide dismutase in sunflower (Helianthus annuus L.)

Polyacrylamide gel electrophoresis of water-soluble proteinsfrom sunflower (Helianthus annuus L.) cotyledons, followed byspecific staining for superoxide dismutase activity, discriminated,according to their electrophoretic mobility, two distinct achromaticbands for Cu/Zn superoxide dismutase. Zymograms of proteinsfrom isolated chloroplasts showed that the chloroplast-locatedCu/Zn superoxide dismutase (Cu/ZnSOD_Chl) migrated faster inthe SOD activity-stained gels. An electrophoretic variant pattern,whose mobility is lower than the control pattern, was identifiedin the ABA-deficient mutant w-1. The variant is coded by a nucleargene with two codominant alleles. Key words: Sunflower, Helianthus annuus L., ABA-deficient mutant, electrophoretic isozyme variant, superoxide dismutase     

Pollen Development and Quantitative Cytochemistry of Exine and Intine Enzymes in Sunflower, Helianthus annuus L

High resolution cytochemical methods have been used to characterizepollen development and pollen-wall structure in Helianthus annuus.Aniline-blue fluorescent material, presumably callose, was detectedin the nexine layer throughout its development. It was associatedwith acid phosphatase activity, while the outer sexine possessedintense esterase activity during the young spore period. Acidphosphatase and esterase were present in both intine and exinewall sites in contrast to their specific location in other pollentypes. Quantitative cytochemical estimates of enzyme activityduring development reveal esterase patterns typical of gametophyticsynthesis, while acid phosphatase patterns are characteristicof sporophytic origin suggesting tapetal transfer during thevacuolate period. Helianthus annuus L, sunflower, pollen development, exine enzymes, intine enzymes, quantitative cytochemistry     

Membrane-bound Plastid Inclusions and Chloroplast Thylakoid Formation in Sunflower (Helianthus annuus L.)

The ultrastructure, distribution and frequency of membrane-boundplastid inclusions present in the epidermal cells of leavesof intact sunflower plants (Helianthus annuus L.) and in theepidermal and mesophyll cells of sunflower leaf discs culturedin darkness have been studied. These inclusions appear to bedilated thylakoids containing a granular material which, undernormal conditions, is probably involved in chloroplast membraneformation. It is suggested that this material accumulates, andinclusions form, in the chloroplasts of sunflower leaves intwo specific situations. Firstly, in the completely differentiatedcells of the epidermis where the chloroplasts, although at arelatively immature stage, have nevertheless reached a terminalstage of development. Secondly, in the mesophyll cells of youngleaves when chloroplast development has been arrested at animmature stage by a 5-day dark period. In the latter situationthe material can be remobilized if plastid development is restimulated.The plastids of sunflower leaf discs cultured in darkness containboth membrane-bound inclusions and prolamellar bodies, indicatingthat they are separate and distinct structures possibly containingdifferent membrane components. Helianthus annuus L., sunflower, chloroplast, ultrastructure, plastid inclusions, thylakoid formation