Phytochromes differentially regulate seed germination responses to light quality and temperature cues during seed maturation

The ratio of red to far-red light (R : FR) experienced by seeds during maturation affects germination, but the genetic regulation of this effect is poorly understood. In Arabidopsis thaliana , responses to R : FR are governed by five phytochrome photoreceptors, PHYA–PHYE . PHYA , PHYB and PHYE mediate germination, but their roles in germination response to the seed maturation environment are largely unknown. Seeds of A. thaliana phytochrome mutants and natural accessions were matured in a factorial combination of cold (16 °C) and warm (24 °C) temperatures and high (R : FR = 1) and low (R : FR = 0.6) R : FR environments, resembling sunlight and foliar shade, respectively. Germination was observed in resulting seeds. All five phytochromes mediated germination responses to seed maturation temperature and/or R : FR environment. PHYA suppressed germination in seeds matured under cold temperature, and PHYB promoted germination under the same conditions. PHYD and PHYE promoted germination of seeds matured under warm temperature, but this effect diminished when seeds matured under reduced R : FR. The A. thaliana natural accessions exhibited interesting variation in germination responses to the experimental conditions. Our results suggest that the role of individual PHY loci in regulating plant responses to R : FR varies depending on temperature and provide novel insights into the genetic basis of maternal effects.     

Illuminated behaviour: phytochrome as a key regulator of light foraging and plant anti-herbivore defence

In many ecological scenarios, the success of an individual plant is defined by the behavioural decisions that it makes when confronted with the risks of competition with other plants, and biomass losses to insect herbivores. These decisions involve expression of shade avoidance responses and induced chemical defences. Because these responses are costly, they frequently engender resource allocation dilemmas. In this review, I discuss the mechanisms that trigger adaptive responses to competitors and herbivores, highlighting the role of phytochromes as central organizers of the overall resource allocation strategy of plants. Phytochromes sense the reduction in the red to far-red (R : FR) ratio of sunlight caused by the proximity of other plants. Shade-intolerant plants respond to low R : FR ratios with shade avoidance behaviours and reduced investment in defence. Pfr depletion leads to increased stability of growth-promoting phytochrome-interacting factors (PIFs), and results in the production of auxins and gibberellins, degradation of DELLA proteins, which are repressors of PIFs, and reduced sensitivity to jasmonates. Thus, phytochrome appears to fulfil its organizational role by regulating the relative strength of the signalling circuits controlled by growth-related and defence-related hormones. I point out cases of signalling redundancy and discuss the significance of recent work on hormone signalling for our understanding of the mechanisms that control adaptive plant behaviour.     

Far-red light reflected from neighbouring vegetation promotes shoot elongation and accelerates flowering in spring barley plants

Field experiments were conducted in St Paul, MN, USA, to test the hypothesis that early season declines in the red:far-red ratio (R:FR) associated with FR reflection from neighbouring leaves have a role in regulating barley development. Treatment plants were grown adjacent to densely sown border rows of barley. The borders functioned to reflect far-red (FR), which reduced R:FR within the treatment plant light environment without shading treatment plants. Barriers were set in the soil to minimize root interactions between treatment plants and borders. Treatment plants were spaced either 2 or 16 cm apart. The presence of borders significantly increased shoot leaf and internode lengths at both plant spacings. Leaf sheath length data suggest that interactions between 2 cm spaced treatment plants enhanced plant responsiveness to the presence of borders. Border treatments shortened the period of vegetative growth prior to initiation of main shoot floral primordia. Bordered plants formed fewer main shoot leaves, initiated internode elongation at a lower node, and had slightly earlier heading dates than unbordered controls. Leaf appearance rate was not influenced by border treatments. We conclude that barley shoot development is photomorphogenically modulated by R:FR. Early season shifts in R:FR could have a significant influence on shoot development given that barley has the capacity to detect and developmentally respond to declines in R:FR associated with FR reflection from neighbours.     

Tree seedling canopy responses to conflicting photosensory cues

Light with decreased red:far-red (R:FR) ratios may signal neighbor presence and trigger plant developmental responses. There is some evidence that plant canopies forage towards increased R:FR ratios, but it is unclear to what extent R:FR versus the total amount of photosynthetically active radiation (PAR) influences canopy foraging responses among forest trees. The objective of this study was to examine the relative importance of PAR and R:FR as photosensory cues leading to tree canopy foraging responses. Seedlings of Betula papyrifera Marshall (paper birch) were grown in an experimental garden. Each seedling was germinated and grown in its own shading structure and exposed to two spatially separated light environments, in a factorial design of PAR and R:FR. Plant canopy foraging was evaluated at the end of one growing season in terms of canopy displacement, canopy area, leaf number, direction of stem lean, petiole aspect, and lamina aspect with respect to experimental light treatments. Leaf number and canopy area were greater on the high PAR sides of plants, irrespective of the R:FR treatment. Seedling canopies were displaced towards the direction of high PAR, but this relationship was not significant across all treatments. Petiole aspect was random and showed no significant directedness towards any of the light treatments. Lamina aspect and the direction of stem lean were distributed towards the direction of high PAR, irrespective of the R:FR treatment. Overall, first-year B. papyrifera seedlings used PAR, rather than R:FR ratio, as a photosensory cue for canopy light foraging.     

Understanding the impact of root morphology on overturning mechanisms: a modelling approach

BACKGROUND AND AIMS: The Finite Element Method (FEM) has been used in recent years to simulate overturning processes in trees. This study aimed at using FEM to determine the role of individual roots in tree anchorage with regard to different rooting patterns, and to estimate stress distribution in the soil and roots during overturning. METHODS: The FEM was used to carry out 2-D simulations of tree uprooting in saturated soft clay and loamy sand-like soil. The anchorage model consisted of a root system embedded in a soil block. Two root patterns were used and individual roots removed to determine their contribution to anchorage. KEY RESULTS: In clay-like soil the size of the root-soil plate formed during overturning was defined by the longest roots. Consequently, all other roots localized within this plate had no influence on anchorage strength. In sand-like soil, removing individual root elements altered anchorage resistance. This result was due to a modification of the shape and size of the root-soil plate, as well as the location of the rotation axis. The tap root and deeper roots had more influence on overturning resistance in sand-like soil compared with clay-like soil. Mechanical stresses were higher in the most superficial roots and also in leeward roots in sand-like soil. The relative difference in stresses between the upper and lower sides of lateral roots was sensitive to root insertion angle. Assuming that root eccentricity is a response to mechanical stresses, these results explain why eccentricity differs depending on root architecture. CONCLUSIONS: A simple 2-D Finite Element model was developed to better understand the mechanisms involved during tree overturning. It has been shown how root system morphology and soil mechanical properties can modify the shape of the root plate slip surface as well as the position of the rotation axis, which are major components of tree anchorage.     

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     

Light signals generated by vegetation shade facilitate acclimation to low light in shade-avoider plants

When growing in search for light, plants can experience continuous or occasional shading by other plants. Plant proximity causes a decrease in the ratio of R to far-red light (low R:FR) due to the preferential absorbance of R light and reflection of FR light by photosynthetic tissues of neighboring plants. This signal is often perceived before actual shading causes a reduction in photosynthetically active radiation (low PAR). Here, we investigated how several Brassicaceae species from different habitats respond to low R:FR and low PAR in terms of elongation, photosynthesis, and photoacclimation. Shade-tolerant plants such as hairy bittercress (Cardamine hirsuta) displayed a good adaptation to low PAR but a poor or null response to low R:FR exposure. In contrast, shade-avoider species, such as Arabidopsis (Arabidopsis thaliana), showed a weak photosynthetic performance under low PAR but they strongly elongated when exposed to low R:FR. These responses could be genetically uncoupled. Most interestingly, exposure to low R:FR of shade-avoider (but not shade-tolerant) plants improved their photoacclimation to low PAR by triggering changes in photosynthesis-related gene expression, pigment accumulation, and chloroplast ultrastructure. These results indicate that low R:FR signaling unleashes molecular, metabolic, and developmental responses that allow shade-avoider plants (including most crops) to adjust their photosynthetic capacity in anticipation of eventual shading by nearby plants.

Vegetation proximity light signals inform shade-avoider plants to adjust their photosynthetic capacity in anticipation of eventual shading by nearby plants.     

Structural development and stability of rice Oryza sativa L. var. Nerica 1

The structural development of glasshouse-grown rice Oryza sativa L. var. Nerica 1 was studied in relation to its stability against lodging. The morphology and mechanical properties of both the stem and roots were examined from tillering, 4 weeks after transplantation up to maturity, together with plant weight distribution and anchorage strength. The "factors of safety" against root and stem failure were subsequently calculated throughout development. Rice plants showed similar morphology to wheat, although they possessed around twice as many tillers per plant and 10 times as many coronal roots. The mechanics of anchorage were also similar. The strength and rigidity of individual tillers increased throughout development as the plants grew taller and heavier and were around 15 times greater than in wheat. By contrast, individual root bending strength, the number of roots, and the anchorage strength levelled off earlier, and anchorage strength was only around twice that in wheat. Consequently, while the self-weight safety factor against stem failure was much higher than in wheat, increasing until late on in development from around 30 to 150, the self-weight safety factor against root anchorage failure was similar to wheat, decreasing from around 15 to 5. Consequently, plants subjected to anchorage tests always failed in their root system rather than their shoot system. The results suggest that, in the field, rice plants would be more likely to undergo root lodging than stem lodging, and that breeding efforts to reduce the incidence of lodging should act to strengthen the rather weak coronal roots.     

Root architecture and tree stability

Summary Root anchorage is discussed with a view to determining the optimum use of root material for enhanced stability. Field observations were made on Sitka spruce root systems while lateral forces were applied to the stem with a winch to pull the tree over. Measurements included the applied force, angles of inclination, soil and root movement, timing of the sound of root breakage using buried microphones, weight and shape of the root-soil plate and damage to the roots.Components of anchorage include the dimensions and mass of the root-soil plate levered from the ground by the displaced stem, and tensile strength of roots and soil beneath the plate; root and soil tensile strength and root/soil resistance on the windward perimeter; and on the lee side the stiffness of the hinge at the fulcrum.Strength properties of roots and soil are reviewed. Models devised for landslip are extended to consider behaviour under tension, of roots singly and in groups, and the concept is developed of a critical rooting density at which root/soil resistance exceeds soil strength, giving rise to the characteric root-soil plate on uprooted trees. The lee side part of the root-soil plate acts as a cantilevered beam and determines the distance of the fulcrum from the tree. Physical laws defining the reduced stiffness of beams as a result of subdivision, indicate the importance of the number/size distribution of roots and weakening effects of branching.On the windward side upward movement of the root-soil plate causes sequential breakage of soil and roots. Under an increasing applied load, failure occurs in parts of the soil-root system before the maximum force for uprooting is achieved. A preliminary approach is made to modelling where the changing contributions of the components of anchorage are allowed for throughout the uprooting process.     

Effects of far-red light on the growth,mycorrhizas and mineral nutrition of Scots pine seedlings

The effects of supplementary far-red sidelight on the formation of mycorrhizas and on the accumulation and allocation of dry weight and mineral nutrients were studied in Scots pine (Pinus sylvestris L.) seedlings. Starting one week after germination the seedlings were subjected to two different light quality regimes: control and simulated sparse-canopy conditions (FR+). In the FR+ regime, light reflected by neighbouring plants was simulated by means of supplementary far-red light sources, which reduced the horizontal red/far-red photon ratio (R:FR) without affecting PAR. Seedlings were harvested after three months of treatment. FR+ increased stem height and decreased the total dry weight of seedlings. Dry weight allocation to needles was not affected, whereas dry weight allocation to roots was reduced and that to stems was increased in FR+ treated seedlings. The total number of short root tips and developing mycorrhizas per seedling were lower in FR+ than in control plants. Most short roots were developing mycorrhizas, while non-mycorrhizal short roots and mycorrhizas with mantle or external mycelium were very scarce. Changes in the allocation of nutrients in general followed the changes in dry weight allocation, and changes in nutrient content followed those in total dry weight. However, mismatches among these changes resulted in significant changes in nutrient concentrations in some organs: the concentrations of nitrogen and potassium in needles and the concentration of nitrogen in stems were higher in FR+ than in control seedlings. Changes in biomass and nutrient allocation under low R:FR may promote rapid height growth during early development in stands of Scots pine seedlings, but concomitant reductions in growth of the root system and mycorrhizas may negatively affect tree performance over the long term.     

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     

The effects of shading and N status on root proliferation in nutrient patches by the perennial grass Agropyron desertorum in the field

Competition for light can affect exploitation of spatially heterogeneous soil resources. To evaluate the influence of shoot status on root growth responses in nutrient-rich soil patches, we studied the effects of shading and whole-plant nitrogen status on root growth in N-enriched and nonenriched patches by mature Agropyron desertorum plants growing in the field with below-ground competition. Roots in enriched patches had greater length to weight ratios (specific root length, SRL), indicating increased absorptive surface areas, compared with roots in control patches. Increased SRL was due to increased production and length of higher order laterals rather than morphological changes in roots of the same branching order. Although the pattern of root growth rates in patches was the same for shaded and unshaded plants, the magnitude of this response to enriched patches was damped by shading. Root relative growth rates (RGR) in N-enriched patches were reduced by more than 50% by short-term shading treatments (60% reduction in photosynthetic flux density), while root RGR in unenriched patches was unaffected by shading. Unexpectedly, plants with higher nitrogen status had greater root RGR in enriched patches than plants that had not received nitrogen supplement, again with no detectable effect on root RGR in the unenriched patches. Therefore, while both shading and plant N status affected the ability of roots to exploit enriched patches by proliferation, there was no stimulation or suppression of root growth in the unenriched, control patches. Thus, plants already under competitive pressure above ground for light and below ground for nutrients should be less able to rapidly respond to opportunities presented in nutrient patches and pulses.     

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     

Branch and root formation in Trifolium repens is influenced by the light environment of unfolded leaves

In plagiotropic plants, axillary buds on the stolon can be exposed to low red:far-red (R:FR) ratios, while the leaves may be positioned in the uppermost layer of the sward where they are exposed to a high R:FR ratio. We tested whether the light environment of unfolded leaves influences outgrowth of the axillary buds and the formation of nodal roots of Trifolium repens. Single plants were grown in a growth cabinet with high photosynthetic photon flux rate (PPFR) and a high R:FR ratio (F_HR_H, control), low PPFR and high R:FR (F_LR_H) or low PPFR and low R:FR (F_LR_L). In an additional treatment (SS), only stolons were shaded so that developing leaves grew into light conditions similar to the control treatment. Neutral shading (F_LR_H) had a minor effect on branching and did not influence root formation. A reduction in the R:FR ratio (F_LR_L) significantly delayed the outgrowth of axillary buds so that, compared to the control plants, the percentage of branched phytomers was reduced by 43% on the parent axis and by 75% on primary branches. Furthermore, the number of nodal roots per plant was reduced by about 30%. When only the stolons were shaded (SS), the percentage of branched and rooted phytomers was similar to that of the control plants. Extension of petioles and leaves was very variable, increasing the values in the F_LR_L treatment at least 2.5-fold compared with the control plants. It was concluded that the light environment of the unfolded leaves had a significant influence on the regulation of the outgrowth of axillary buds and that the high plasticity in petiole growth allows the positioning of the leaves in a light environment conducive to the stimulation of branch outgrowth. Received: 8 February 1997 / Accepted: 26 April 1997     

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     

The interaction of light quality and irradiance with gibberellins, cytokinins and auxin in regulating growth of Helianthus annuus hypocotyls

A reduced red to far-red (R/FR) light ratio and low photosynthetically active radiation (PAR) irradiance are both strong signals for inducing etiolation growth of plant stems. Under natural field conditions, plants can be exposed to either a reduced R/FR ratio or lower PAR, or to a combination of both. We used Helianthus annuus L., the sunflower, to study the effect of reduced R/FR ratio, low PAR or their combination on hypocotyl elongation. To accomplish this, we attempted to uncouple light quality from light irradiance as factors controlling hypocotyl elongation. We measured alterations in the levels of endogenous gibberellins (GAs), cytokinins (CKs) and the auxin indole-3-acetic acid (IAA), and the effect of exogenous hormones on hypocotyl growth. As expected, both reduced R/FR ratio and lower PAR can significantly promote sunflower hypocotyl elongation when given separately. However, providing the reduced R/FR ratio at a low PAR resulted in the greatest hypocotyl growth, and this was accompanied by significantly higher levels of endogenous IAA, GA1, GA8, GA20 and of a wide range of CKs. Providing a reduced R/FR ratio under normal PAR also significantly increased growth and again gave significantly higher levels of endogenous IAA, GAs and CKs. However, only under the de-etiolating influence of a normal R/FR ratio did lowering PAR significantly increase levels of GA1, GA8 and GA20. We thus conclude that light quality (e.g. the R/FR ratio) is the most important component of shade for controlling hypocotyl growth and elevated growth hormone content.     

The effect of nitrogen and growth regulators on stem and root characteristics associated with lodging in two cultivars of winter wheat

The effects of nitrogen and plant growth regulators (stem shorteners)on root and shoot characteristics associated with lodging resistancewere investigated in two winter wheat (Triticum aestivum L.)cultivars of contrasting lodging resistance: the susceptibleGalahad and the resistant Hereward. The morphology and mechanicalstrength of the stems and anchorage systems grown at two levelsof nitrogen and with or without growth regulators were measuredand related to the incidence of lodging recorded in a fieldtrial. In both cultivars high levels of nitrogen increased theheight of the stem, thereby increasing the ‘self-weight’moment transmitted into the ground and weakened both the stemsand the anchorage coronal roots. As a result, the anchoragestrength was also reduced, plants failing in the root systemin simulated lodging tests. Growth regulators, in contrast,had little effect on the bending strength of the shoots androot systems, but reduced plant height so that the over turningmoments generated by the weight of the shoot were less. Therewere also differences between cultivars: Galahad plants hadweaker anchorage due to the smaller number and lower strengthof the coronal roots. The morphological and mechanical measureswere used to calculate a safety factor against both stem androot lodging. Five factors were found to influence the safetyfactors, these were: cultivar type, the type of lodging, therate of nitrogen and growth regulator application, and time,being lowest in Galahad plants at high levels of nitrogen andwithout growth regulators and at grain filling when the earswere heaviest. This was consistent with the observed patternof lodging: root lodging occurred at grain filling and onlyin Galahad which had been treated with high nitrogen rates,most strongly in plants without growth regulators. Key words: Lodging, safety factors, anchorage, ‘self-weight’ moment     

The influence of slope on <Emphasis Type="Italic">Spartium junceum</Emphasis> root system: morphological,anatomical and biomechanical adaptation

Root systems have a pivotal role in plant anchorage and their mechanical interactions with the soil may contribute to soil reinforcement and stabilization of slide-prone slopes. In order to understand the responses of root system to mechanical stress induced by slope, samples of Spartium junceum L., growing in slope and in plane natural conditions, were compared in their morphology, biomechanical properties and anatomical features. Soils sampled in slope and plane revealed similar characteristics, with the exception of organic matter content and penetrometer resistance, both higher in slope. Slope significantly influenced root morphology and in particular the distribution of lateral roots along the soil depth. Indeed, first-order lateral roots of plants growing on slope condition showed an asymmetric distribution between up- and down-slope. Contrarily, this asymmetric distribution was not observed in plants growing in plane. The tensile strength was higher in lateral roots growing up-slope and in plane conditions than in those growing down-slope. Anatomical investigations revealed that, while roots grown up-slope had higher area covered by xylem fibers, the ratio of xylem and phloem fibers to root diameter did not differ among the three conditions, as also, no differences were found for xylem fiber cell wall thickness. Roots growing up-slope were the main contributors to anchorage properties, which included higher strength and higher number of fibers in the xylematic tissues. Results suggested that a combination of root-specific morphological, anatomical and biomechanical traits, determines anchorage functions in slope conditions.     

Far-red enrichment and photosynthetically active radiation level influence leaf senescence in field-grown sunflower

Basal leaves frequently senesce before anthesis in high population density crops. This paper evaluates the hypothesis that quantitative and qualitative changes in the light environment associated with a high leaf area index (LAI) trigger leaf senescence in sunflower ( Helianthus annuus L.) canopies. Mean leaf duration (LD, time from achievement of maximum leaf area) of leaf 8 was significantly ( P < 0.05) reduced from 51 to 19 days as crop population density was increased from 0.47 to 4.76 plants m⁻². High compared to low plant population density was associated with earlier reduction in the photosynthetically active radiation (PAR) and red/far-red ratio (R/FR) reaching the target leaf. However the changes in R/FR preceded those in PAR. When the light environment of individual leaves of isolated plants growing under field conditions was manipulated using filters and FR-reflecting mirrors, LD was positively and linearly related with the mean daily PAR (MDR) received in the FR- (no FR enrichment) treatments. FR enrichment of light reaching the abaxial surface of the leaf significantly ( P < 0.05) reduced LD by 9 days at intermediate PAR levels with respect to FR-controls, but did not affect LD at the maximum PAR used in these experiments. However, when light reaching both leaf surfaces was enriched with FR, LD (for leaves receiving maximum PAR) was 13 days shorter than that of the FR- control. These results show that basal leaf senescence in sunflower is enhanced both by a decrease in PAR and by a decrease in R/FR.