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     

Responses of the Root Systems of Sunflower and Maize to Unidirectional Stem Flexure

Plants of two contrasting species of herbaceous annuals, thedicot sunflower (Helianthus annuusL.) and the monocot maize(Zea maysL.), grown in the glasshouse were subjected to regularunidirectional stem flexure. Differences in morphology and mechanicalproperties of roots and shoots were then investigated. Rootsystems were divided into quadrants around the axis of stimulationand differences in root morphology and mechanics between thezones were investigated. There were considerable differencesbetween roots in the leeward and windward zones compared withroots perpendicular to the axis of stimulation. First-orderlateral roots in both species were thicker, more rigid and morenumerous. These results suggest that plant roots respond locallyto mechanical stimulation. There were, however, also differencesin the responses of the two species. In sunflower, the tap rootand stem base became elliptical in cross section with the majoraxis lying in the plane of stimulation. The lateral roots offlexed sunflowers in both the leeward and windward zones showedsimilar growth responses: roots were thicker, more numerousand weighed more than those in the perpendicular zones. However,only leeward roots showed significant differences in their mechanicalproperties; roots were more rigid, stronger and stiffer. Incontrast, the leeward roots of maize were thicker and more numerous,with a greater biomass than the windward roots. However, onlyroots in the windward zone were stiffer than those in the perpendicularzone. These differences between sunflower and maize are relatedto their contrasting anchorage mechanics.Copyright 1998 Annalsof Botany Company Anchorage, biomechanics, adaptive growth, roots, thigmomorphogenesis,Helianthus annuusL.,Zea maysL.     

Involvement of the Cotyledon in Adventitious Root Initiation in Impatiens balsamina L. Seedlings

Adventitious root primordia are found in the pre-hypocotyl tissueof developing seeds of Impatiens balsamina L. by the third weekafter petal drop, and are present in the mature seed. Aftergermination, the adventitious roots emerge from a collet swellingon the hypocotyl of the young seedlings. Removal of the colletduring the first five days results in the formation of anotherat the base of the remaining hypocotyl. Older seedlings respondto the excision of the collet by producing one or more rootsnear the cut end, unless the cut is made close to the cotyledon,when, even in nine-day seedlings, a reduced collet is formedassociated with four or fewer roots. The influence of the cotyledonon collet/root regeneration diminishes in older seedlings andin these is manifested only in hypocotyl tissue adjacent tothat organ. Impatiens balsamina, balsam, cotyledon, adventitious roots, collet     

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 Mechanics of Anchorage in Wheat Triticum aestivum L.: II. ANCHORAGE OF MATURE WHEAT AGAINST LODGING

The root system of mature wheat Triticum aestivum Marts Doveis dominated by the 7 to 15 adventitious roots which emergefrom the perimeter of the stem base, pointing radially outwardsand downwards. The basal, coronal region of these roots is thickand unbranched, attached to a rhizosheath of earth by a densecovering of root hairs and stiffened in bending by lignificationof outer layers of the cortex. Root lodging of plants involves bending of the coronal rootsat their base and axial movement of leeward and windward rootsthrough the soil; their resistance to these motions providemoments resisting lodging. A model of anchorage was producedby summing the resistance of each root to both forms of motionto give two anchorage components. The model was tested in aseries of mechanical experiments in which simulated lodgingwas followed by loading of individual roots; results supportedthe anchorage model and suggested that in the experimental conditionsthe two components of anchorage were approximately equal inmagnitude. The stem was about 30% stronger than the anchoragesystem. The coronal anchorage roots made up 4.4% of total dry mass;it is suggested that anchorage could be improved either by increasinginvestment in this region or by altering root orientation. Sequentialdevelopment of seminal and adventitious root systems is relatedto the changes in anchorage requirement with age.     

Patterns of Regeneration in Mutilated Seedlings of Garden Balsam, Impatiens balsamina L.

Seedlings of garden balsam, Impatiens balsamina, form an encirclingswollen ring, the collet, basal to the hypocotyl. From this,four lateral roots emerge promptly after germination. When thisstructure is excised, it regenerates as a basal encircling swellingfrom which four lateral roots with root hairs emerge. Repetitionof the removal of this structure results again in the regenerationof a similar complex. Eventually, after 3–4 excisions,the pattern is broken and lateral roots occur sporadically alongthe hypocotyl. Mutilations of seedlings of the garden balsamindicate that the regeneration will not occur in the absenceof cotyledonary tissue. This suggests a control site for thisregenerative phenomenon but no mechanism for this control isadvanced. Impatiens balsamina, balsam, lateral roots, regeneration, collet     

The function of buttress roots: a comparative study of the anchorage systems of buttressed (Aglaia and Nephelium ramboutan species) and non-buttressed (Mallotus wrayi) tropical trees

The anchorage mechanics of mature buttressed trees of Aglaiaand Nephelium, and of non-buttressed Mallotus wrayi have beeninvestigated by combining a study of the morphology of theirroot systems with a series of anchorage tests. Both types possessed tap roots, but only buttressed trees possessedsinker roots, which branched from the ends of the buttresses.The anchorage strength of the buttressed trees was almost double(10.6 kNm) that of the unbuttressed ones (4.9 kNm), and themaximum moment was generated at lower angles. In but tressedtrees, the leeward buttresses were pushed into the soil beforebending and eventually breaking towards their tip, whilst thewindward buttresses pulled out of the soil or delaminated ifthey possessed sinker roots. The tap root rotated in the soilto windward. In contrast, during failure of unbuttressed treesthe tap root both moved and bent towards the leeward, the windwardroots were pulled out of the soil, and the leeward lateralssimply buckled. Strains along but tresses were much higher thanalong the laterals of unbuttressed trees. These results suggest that buttresses act in both tension andcompression and make a much larger contribution to anchoragethan the thin laterals of non-buttressed trees. The relativecontribution of the but tresses was determined by carrying outa further series of anchorage tests in which both buttressedand unbuttressed trees were pulled over after all their lateralshad been cut away. These trees were therefore only anchoredby their taproot. Failure of both types was similar to intactunbuttressed trees, and they had similar anchorage strengthstoeach other, 4 kNm, around 80% of the value for intact non-buttressedtrees, but only 40% of the strength of intact buttressed trees.Buttresses therefore contribute around 60% of the anchorageof buttressed trees, producing around six times more anchoragethan the thin laterals of unbuttressed trees. Key words: Anchorage, root architecture, sinker roots, tap roots, root bending strength, buttresses     

Factors Influencing Adventitious Root Production in Cuttings of Griselinia littoralis and Griselinia lucida

Griselinia littoralis roots quickly and vigorously from cuttingsof seedlings and mature plants and also forms roots on detachedleaves. Cuttings root in the dark but leaves must be present.In contrast G. lucida roots vigorously only from cuttings takenfrom seedlings. Light is essential for root formation on oldermaterial. Detached leaves will not root. Although callus formationat the cut base commonly occurs in both species it is not directlyassociated with root formation. Griselinia littoralis Raoul Choix, Griselinia lucida Forst. f. Prodr., adventitious roots, stem cuttings, leaf cuttings, woody plants     

The anchorage mechanics of deep rooted larch, Larix europea L. japonica

The anchorage of deep rooted 16-year-old larch trees, Larixeuropea japonica, has been studied by combining winching testswith analyses of strain around the base of the trunk and rootsystem and mechanical tests on individual roots. These showedthat anchorage is provided by the laterals which emerge fromaround the stem base, sinker roots which emerge along theirlength, and tap roots positioned directly underneath the bole.During anchorage failure the leeward laterals are bent and eventuallybreak close to their base, whilst the windward laterals arepulled out of the ground, with their sinker roots intact. Afterinitially being confined by the soil and bending, the tap rootrotates in the soil. Anchorage failure is similar when the soilis dry as when it is wet, but failure occurs closer to the trunk.Strain measurements along the lateral roots revealed that thestresses were highest close to the trunk and that these regionsof the roots contribute most to tree stability. The two major components of anchorage were found to be the resistanceof leeward laterals to bending and the resistance of tap rootsand windward sinkers to uprooting. Bending tests on leewardlaterals revealed that they provide around 25% of tree anchorage.Almost 75% of the anchorage strength must, therefore, be providedby the windward sinkers and tap roots. Anchorage strength ofroots was positively correlated to their cross-sectional area.The vertical orientation of the sinkers makes the anchoragesystem of larch more efficient than the plate system formedby Sitka spruce on waterlogged soils and means that no root-soilplate is formed. Key words: Anchorage, root architecture, sinker roots, root bending strength, windthrow     

Adventitious Root Formation in Veronica Spp.

Species of the genus Veronica differ in habitat preferences,growth form and in adventitious root production. The annualspecies rarely or never produce adventitious roots in intactplants in the field but some, for example V. persica and V.arvensis will root vigorously from single node stem segmentsin culture. Others, such as V. agrestis require the presenceof IAA for substantial levels of root formation to occur incultured stem segments. Veronica hederifolia cuttings rarelyproduce roots. Stem cuttings of the perennial species, in general,rooted more vigorously than those of annual plants. Both V.fihiformis and V. serpyllifolia root very strongly. The position of root production from the stem cuttings differedfrom species to species. Roots arose either from the node, theregion of the base or at some intermediate point. Veronica arvensis,V. chamaedrys and V. persica rooted mainly from the basal regionwhereas V. filiformis rooted mainly from the node. Veronicaserpyllifolia cuttings rooted at both of these locations. Veronica filiformis, a perennial species that is infertile inBritain, produces root primordia in intact plants at nodes whichare close to the shoot apex. Thus, even very young stem segmentshave ‘preformed’ root primordia. For this reason,detached stem segments of V. filiformis root very rapidly andthis probably has been of great significance in its successfulinvasion and spread in lawns and short turf areas. Veronica spp., adventitious roots, indol-3-ylacetic acid, root primordia, vegetative reproduction     

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     

Gravitropism of Primary Roots of Zea mays L. at Different Displacement Angles

Primary roots of Zea mays were oriented at various angles fromthe vertical ranging from 99° to 1° and their subsequentbending analysed from filmed records. The maximum rate of bendingand the time before bending commenced both varied two-fold,but showed no correlation with the initial angle of tip displacement.Roots orientated to small initial angles (< 40°) oftenovershot the vertical and proceeded to oscillate around thisorientation, whereas roots oriented to large initial angles(> 60°) often failed to achieve the vertical. Roots inthis latter group resumed bending after an indeterminate time,or did so immediately after a second displacement of their tip,showing that they were not intrinsically unable to bend. Theapparently spontaneous resumption of bending after a temporaryplagiogravitropic phase is suggested as being due to noise inthe graviperception system in the root cap. The tips of rootsgrowing vertically downwards showed oscillatory bending movementsup to 10° either side of vertical. This angle correspondsto the minimum angle of displacement which induces gravitropicbending. Only when roots were oriented 10-20° from verticaldid they begin unequivocally to show a gravitropism since atsuch angles the deflection of their tips exceeded that due totheir natural oscillation.Copyright 1993, 1999 Academic Press Gravitropism, roots, Zea mays     

Anatomy and Histochemistry of the Root System of the Kiwifruit Vine, Actinidia deliciosa var. deliciosa.

The kiwifruit vine is a species which has been newly introducedinto cultivation and little is known of its comparative physiologyand anatomy. In this study we found that fibrous, 'magnolioid'roots, which have undergone secondary vascular development butwhich retain the cortex and develop a suberized epidermis, comprisethe greater part of the root system (95% of total length). Newlyinitiated roots with primary development conform to norms establishedin other woody plant species. However, the structural roots,like the fibrous roots, also retain a cortex and phellodermwhich is initiated by hypodermal cells within the cortex andnot by the pericycle which is the common progenitor tissue inother species. This phellogen produces new cells centrifugallyonly. The cortex is a relatively small component of the structuralroot and the bulk of the tissue is vascular in origin, as inthe roots of other plant species. The endodermis is retainedand continues to divide periclinally to accommodate the increasein circumference with growth.Copyright 1993, 1999 Academic Press Actinidia deliciosa, root anatomy, ontogony, histochemistry, exodermis, endodermis     

The Mechanics of Root Lodging in Winter Wheat, Triticum aestivum L.

The anchorage of winter wheat, Triticum aestivum L., is providedby a cone of rigid coronal roots which emerge from around thestem base. During root lodging this cone rotates at its windwardedge below the soil surface, the soil inside the cone movingas a block and compressing the soil beneath. A theoretical modelof anchorage suggested that lodging resistance should be dependenton the diameter of the root-soil cone, coronal root bendingstrength and soil shear strength. We tested the predictions of the anchorage model by carryingout two series of experiments. In the first, varieties of contrastinglodging resistances were artificially lodged. The moment requiredto rotate plants into the soil, the diameter of the root-soilcone, and the bending strength of the coronal roots were recorded.The lodging moment was correlated with the size of the soilcone, as predicted. Generally, differences in anchorage strengthbetween varieties were due to differences in root-soil conediameter, although coronal root strength was also important. A second series of tests was carried out using model plantsanchored by plastic discs. The behaviour of the models duringartificial lodging supported the anchorage model; the forceresisting lodging was similar to that of plants with root-soilcones of the same size and the resisting force was dependenton the soil strength. These results suggest that root lodging resistance might beimproved by increasing both the angle of spread and the bendingstrength of the coronal roots. Key words: Anchorage, root-soil cone, coronal roots, lodging, wheat     

A Relationship between Phosphate Uptake and Membrane Electrical Potential in Excised Roots of Helianthus annuus

Phosphate uptake by excised roots of sunflower (Helianthus annuus)was determined by the disappearance of phosphate from the externalsolution and by the accumulation of phosphate labelled with³²P. Over a 24 h period it was observed that net phosphate uptakedeclined to zero whilst uptake of ³²P continued unabated. Theelectrical PD of the cortical cell membranes declined in parallelwith net phosphate uptake and it was found that both could berestored by creating a pH gradient across the plasmalemma. Itwas concluded that net phosphate uptake was responsible fora component of the membrane PD of the root cortical cells. Key words: Roots, Phosphate, Membranes     

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.     

Strain distribution during anchorage failure of Pinus pinaster Ait. at different ages and tree growth response to wind-induced root movement

Winching tests were carried out on 5- 13- and 17-year-old tap rooted Maritime pine (Pinus pinaster Ait.) in order to determine how the mode of anchorage failure changes throughout the life of a tree. As trees were pulled sideways, strain along the lateral roots was recorded using strain gauges attached to a strain indicator. Measurements of strain in the root system, taken during winching, provide information about root movement when loaded by wind. The mode of mechanical failure appeared to depend on tree age. The youngest trees bent over completely during winching, but did not break due to the plasticity of their trunks. The 13-year-old trees either broke at the base of the tree (due to the presence of grafting scar tissue) or at the base of the tap-root. The oldest trees broke at the base of the tap-root and sounds of roots breaking were also heard. Strain was twice as great in the trunk compared to the roots in the 5- and 13-year-old trees and was three times greater in the compression roots of 17-year–old trees compared to that in the trunk. In one 17-year-old tree, strain was found to increase at a distance of 35 cm in tension roots before decreasing again. Although the mode of failure changed with tree size, anchorage strength increased proportionally with the third power of trunk diameter, therefore another reason why failure differs with tree age must exist. In order to determine if different types of wood were being laid down in the lateral roots in response to wind loading, maturation strains, indicating the existence of mechanical stress in developing wood cells, were measured at different points along the roots. A high correlation was found between maturation strain and strain measured during winching, in roots that lay in the wind direction only. Therefore, trees appear to be able to respond to external loading stress, even at a local level within a root. This revised version was published online in June 2006 with corrections to the Cover Date.     

The Mechanics of Anchorage in Wheat Triticum aestivum L.: I. THE ANCHORAGE OF WHEAT SEEDLINGS

Model of the mechanics of uprooting lead to the identificationof ‘optimal’ anchorage systems which can withstanda given upward force at a minimum construction cost. Such systemshave many downward-pointing fibrous roots which are strengthenedprogressively towards the base. A study of the anchorage systemof 7- and 21-d-old wheat (Triticum aestivum L.) plants showedthat the plants possessed five seminal roots, of which onlythree pointed vertically. Each root was well suited for anchorage,being convered in root hairs and strengthened progressivelytowards the base by lignification of the stele. Strength andstiffiness of roots but not their mass per unit length increasedwith age. There was little interaction between roots when plantswere uprooted; the three vertical roots broke while the twohorizontal ones pulled out, as occurred when roots were pulledout singly, Uprooting forces increased with age and the rootsystem could withstand uprooting forces greater than those requiredto pull out upper leaves, so reducing the chances of the plantbeing uprooted by a herbivore, By 3 weeks a stiff adventitiousroot system, which would later help prevent the wheat lodging,was developing.     

Comparative Root Anatomy of Five Actinidia Species in Relation to Rootstock Effects on Kiwifruit Flowering

This study examined the root anatomy of five Actinidia specieswhich, when used as rootstocks, differ in their effects on flowerproduction of the 'Hayward' kiwifruit scion. The rootstock speciesexamined were A. hemsleyana, A. eriantha, A. rufa, A. deliciosaand A. chinensis. Roots of all the five species had similartissues and similar cell types in each tissue. The cortex andendodermis were retained and developed further during secondaryroot thickening. Each meriphyte of endodermal cells, derivedduring secondary growth, was enclosed in a suberin lamella.Crystalline idioblasts were present in the cortex and phloem.These idioblasts contained raphide crystals, often surroundedby mucilage. Roots of the five species differed significantlyin the total cross-sectional area of vessels in the xylem, inthe total cross-sectional areas of fibres and crystalline idioblastsin the phloem, and in the abundance of starch grains in parenchymaof all tissues. Roots of flower-promoting rootstock speciestended to have more and larger xylem vessels, more crystallineidioblasts, and more starch grains. These anatomical featuresmay be used as criteria for future selection of flower-promotingrootstocks in kiwifruit. The possible physiological link betweenroot anatomy and rootstock effect is discussed.Copyright 1994,1999 Academic Press Kiwifruit, Actinidia, root, anatomy, rootstock, flowering     

The Influence of Mineral Nutrition on the Root Development of Trees: I. THE GROWTH OF SITKA SPRUCE WITH DIVIDED ROOT SYSTEMS

Sitka spruce seedlings were grown with their root systems dividedbetween two contrasting nutrient regimes. One half of the rootsystem was supplied with a solution containing N, P, and K ata range of concentrations while the untreated half receivedwater only. High-nutrient treatments induced two flushes ofshoot growth resulting in a large shoot system, whereas plantsin the low-nutrient treatments flushed once only and showedsymptoms of nutrient deficiency. Root growth, assessed in terms of dry weight and diameter ofboth primary and woody tissues, was stimulated in the rootsto which the nutrients were actually applied, whereas in theuntreated roots on treated plants only the primary root diameterwas enhanced. However, internal nutrient concentrations on bothsides of the root system were related to the concentrationsapplied, though to a slightly lesser extent in the untreatedroots. Thus, the nutrients which had been internally translocatedto the untreated roots had little effect on their growth. The localized stimulation of xylem production in the woody rootsextended into the stem along a spiral pathway which was demonstratedby the movement of dye. Possible mechanisms are discussed by which differential rootgrowth is brought about by a localized supply of mineral nutrients.