How kelp in drag lose their ruffles: environmental cues, growth kinematics, and mechanical constraints govern curvature.

We reveal how patterns of growth in response to environmental cues can produce curvature in biological structures by setting up mechanical stresses that cause elastic buckling. Nereocystis luetkeana are nearshore kelp with wide ruffled blades that minimize self-shading in slow flow, but narrow flat blades that reduce hydrodynamic drag in rapid flow. Previously we showed that blade ruffling is a plastic trait associated with a transverse gradient in longitudinal growth. Here we consider expansion and displacement of tissue elements due to growth in blades, and find that growth patterns are altered by tensile stress due to hydrodynamic drag, but not by shading or nutrients. When longitudinal stress in a blade is low in slow flow, blade edges grow faster than the midline in young tissue near the blade base. Tissue elements are displaced distally by expansion of younger proximal tissue. Strain energy caused by the transverse gradient in longitudinal growth is released by elastic buckling once the blade grows wide enough, producing ruffles distal to the region where the growth inhomogeneity started. If a blade experiences higher stress in rapid flow, the edges and midline grow at the same rate, so the blade becomes flat as these new tissue elements are displaced distally.

The role of root development of Avena fatua in conferring soil strength.

UNLABELLED: • PREMISE OF THE STUDY: Roots play an important role in strengthening and stabilizing soils. Existing models predict that tensile strength and root abundance are primary factors that strengthen soil. This study quantified how both factors are affected by root developmental stage.• METHODS: Focusing on early development of Avena fatua, a common grassland species with a fibrous root system, we chose three developmental stages associated with major changes in the root system. Seeds were planted in rhizotrons for easy viewing and pots to allow root growth surrounded by soil. Tensile strength was determined by subjecting root segments to a progressively larger pulling force until breaking occurred. Root abundance at two depths was characterized by the cross-sectional area of the roots divided by the area of the soil core (i.e., root area ratio). Shear strength of 50 mm saturated soil columns was determined with a modified interface direct shear device.• KEY RESULTS: Tensile strength increased by a factor of ≥15× with distance from the root tip. Thus, soil-strengthening properties increased with root cell development. Plants grown under dry soil conditions produced roots with higher maximal tensile strength (41.9 MPa vs. approximately 17 MPa), largely explained by 33% thinner diameters. Over 7 weeks of root growth, root abundance increased by a factor of 4.8× while saturated soil shear strength increased by 24% in the upper soil layer.• CONCLUSIONS: Root development should be incorporated into models of soil stability to improve understanding of this important environmental property.

Deposition of ammonium and nitrate in the roots of maize seedlings supplied with different nitrogen salts.

This study measured total osmolarity and concentrations of NH(4)(+), NO(3)(-), K(+), soluble carbohydrates, and organic acids in maize seminal roots as a function of distance from the apex, and NH(4)(+) and NO(3)(-) in xylem sap for plants receiving NH(4)(+) or NO(3)(-) as a sole N-source, NH(4)(+) plus NO(3)(-), or no nitrogen at all. The disparity between net deposition rates and net exogenous influx of NH(4)(+) indicated that growing cells imported NH(4)(+) from more mature tissue, whereas more mature root tissues assimilated or translocated a portion of the NH(4)(+) absorbed. Net root NO(3)(-) influx under Ca(NO(3))(2) nutrition was adequate to account for pools found in the growth zone and provided twice as much as was deposited locally throughout the non-growing tissue. In contrast, net root NO(3)(-) influx under NH(4)NO(3) was less than the local deposition rate in the growth zone, indicating that additional NO(3)(-) was imported or metabolically produced. The profile of NO(3)(-) deposition rate in the growth zone, however, was similar for the plants receiving Ca(NO(3))(2) or NH(4)NO(3). These results suggest that NO(3)(-) may serve a major role as an osmoticant for supporting root elongation in the basal part of the growth zone and maintaining root function in the young mature tissues.

To duckweeds (Landoltia punctata), nanoparticulate copper oxide is more inhibitory than the soluble copper in the bulk solution.

CuO nanoparticles (CuO-NP) were synthesized in a hydrogen diffusion flame. Particle size and morphology were characterized using scanning mobility particle sizing, Brunauer-Emmett-Teller analysis, dynamic light scattering, and transmission electron microscopy. The solubility of CuO-NP varied with both pH and presence of other ions. CuO-NP and comparable doses of soluble Cu were applied to duckweeds, Landoltia punctata. Growth was inhibited 50% by either 0.6 mg L(-1) soluble copper or by 1.0 mg L(-1) CuO-NP that released only 0.16 mg L(-1) soluble Cu into growth medium. A significant decrease of chlorophyll was observed in plants stressed by 1.0 mg L(-1) CuO-NP, but not in the comparable 0.2 mg L(-1) soluble Cu treatment. The Cu content of fronds exposed to CuO-NP is four times higher than in fronds exposed to an equivalent dose of soluble copper, and this is enough to explain the inhibitory effects on growth and chlorophyll content.

Environmental effects on spatial and temporal patterns of leaf and root growth.

Leaves and roots live in dramatically different habitats, but are parts of the same organism. Automated image processing of time-lapse records of these organs has led to understanding of spatial and temporal patterns of growth on time scales from minutes to weeks. Growth zones in roots and leaves show distinct patterns during a diel cycle (24 h period). In dicot leaves under nonstressful conditions these patterns are characterized by endogenous rhythms, sometimes superimposed upon morphogenesis driven by environmental variation. In roots and monocot leaves the growth patterns depend more strongly on environmental fluctuations. Because the impact of spatial variations and temporal fluctuations of above- and belowground environmental parameters must be processed by the plant body as an entire system whose individual modules interact on different levels, growth reactions of individual modules are often highly nonlinear. A mechanistic understanding of plant resource use efficiency and performance in a dynamically fluctuating environment therefore requires an accurate analysis of leaf and root growth patterns in conjunction with knowledge of major intraplant communication systems and metabolic pathways.

Modeling the hydraulics of root growth in three dimensions with phloem water sources.

Primary growth is characterized by cell expansion facilitated by water uptake generating hydrostatic (turgor) pressure to inflate the cell, stretching the rigid cell walls. The multiple source theory of root growth hypothesizes that root growth involves transport of water both from the soil surrounding the growth zone and from the mature tissue higher in the root via phloem and protophloem. Here, protophloem water sources are used as boundary conditions in a classical, three-dimensional model of growth-sustaining water potentials in primary roots. The model predicts small radial gradients in water potential, with a significant longitudinal gradient. The results improve the agreement of theory with empirical studies for water potential in the primary growth zone of roots of maize (Zea mays). A sensitivity analysis quantifies the functional importance of apical phloem differentiation in permitting growth and reveals that the presence of phloem water sources makes the growth-sustaining water relations of the root relatively insensitive to changes in root radius and hydraulic conductivity. Adaptation to drought and other environmental stresses is predicted to involve more apical differentiation of phloem and/or higher phloem delivery rates to the growth zone.

Laser ablation ICP-MS reveals patterns of copper differing from zinc in growth zones of cucumber roots.

Laser ablation coupled with inductively coupled plasma-mass spectrometry was used to find Cu and Zn concentration in surface tissue along a longitudinal developmental gradient with meristem, rapidly elongating tissue, and nongrowing tissue in a model system of seedling roots of Cucumis sativus L. (cucumber). Tissue metal accumulation was determined for roots of seedlings growing on cellulosic germination paper treated with nutrient solution (controls), and also treated with concentrations of Zn (40 ppm) and Cu (10 ppm) that reduced growth. Cu content of all roots is highest at the apex and falls sharply to lower values by 2 mm from the root tip. In contrast, at moderate Zn availability (0.07 ppm), Zn content rises from the apex to 2 mm then falls throughout the remainder of the growth zone. At high external Zn the spatial pattern resembles that of Cu. Cucumber root growth zones accumulate more of each metal with higher external availability. Metal deposition rates were calculated using a continuity equation with data on local metal content and growth velocity. Deposition rates of both metals are generally highest in the rapidly elongating region, 1.5-3.5 mm, even where metal concentration is decreasing with position and root age and even when the accumulation is inhibitory to growth.

Moving with climbing plants from Charles Darwin's time into the 21st century.

We provide an overview of research on climbing plants from Charles Darwin to the present day. Following Darwin's interests, this review will focus on functional perspectives including attachment mechanisms and stem structure and function. We draw attention to a number of unsolved problems inviting future research. These include the mechanism for establishment of the twining habit, a quantitative description following the development of a tissue element through space and time, the chemistry of sticky exudates, the microstructure of xylem and the capacity for water storage, the vulnerability to embolism, and the mechanism for embolism repair. In conclusion we cite evidence that, in response to increasing CO(2) concentration, anthropic perturbation and/ or increasing forest fragmentation, lianas are increasing relative to tree species. In the 21st century, we are returning to the multiscale, multidisciplinary approach taken by Darwin to understand natural history.

Stage-dependent border cell and carbon flow from roots to rhizosphere.

Rising CO(2) levels in the atmosphere have drawn attention to the important role of soil in sequestering carbon. This project goal was to quantify soil carbon deposition associated with border cell release and exudation from root growth zones. Carbon was measured with a Carlo Erba C/N analyzer in soil from the rhizosphere of mature grasses and, in separate experiments, in soil collected around root growth zones. Root border cells in "rhizosphere soil" (silica sand) were counted using a compound microscope after soil sonication and extraction with surfactant. For sand-grown Bromus carinatus, Zea mays, and Cucumis sativus, young seedlings (with roots shorter than 2 cm) released thousands of border cells, while older root tips released only hundreds. For a variety of native annual and perennial grasses and invasive annual grasses (Nassella pulchra, B. carinatus, B. diandrus, B. hordeaceus, Vulpia microstachys, Aegilops triuncialis, Lolium multiflorum, Zea mays), the rhizosphere of mature root systems contained between 18 and 32 µg C g(-1) sand more than that of the unplanted controls. Spatial analysis of the rhizosphere around the cucumber growth zone confirmed C enrichment there. The root tip provided C to the rhizosphere: 4.6 µg C in front of the growing tip, with the largest deposition, 20.4 µg C, to the rhizosphere surrounding the apical 3 mm (root cap/meristem). These numbers from laboratory studies represent the maximum C that might be released during flooding in soils. Scaling up from the organ scale to the field requires a growth analysis to quantify root tip distributions in space and time.

Compost amendment of Cu-Zn minespoil reduces toxic bioavailable heavy metal concentrations and promotes establishment and biomass production of Bromus carinatus (Hook and Arn.).

A series of lab and greenhouse studies were undertaken to understand how Cu and Zn toxicity influences Bromus carinatus (Hook and Arn.) growth, to what degree an organic amendment (yard waste compost) may reduce Cu and Zn bioavailability in Cu-Zn minespoil and promote plant growth in combination with fertilizer, and how the vertical distribution of compost in the minespoil influences rooting depth. Root Cu and Zn toxicity thresholds were determined to be 1 mgL(-1) and 10 mgL(-1) in solution, respectively. The compost amendment had exceptionally high Cu and Zn binding capacities (0.17 and 0.08 g metal g C(-1), for Cu and Zn, respectively) that were attributed to high compost humic and fulvic acid concentrations. Maximum plant biomass was achieved when minespoil was amended with compost and fertilizer in combination. Fertilizer alone had no effect on plant growth. Mixing compost into the minespoil was essential to promote adequate rooting depth.

Seasonal and spatial patterns of metals at a restored copper mine site. I. Stream copper and zinc.

Seasonal and spatial variations in metal concentrations and pH were found in a stream at a restored copper mine site located near a massive sulfide deposit in the Foothill copper-zinc belt of the Sierra Nevada, California. At the mouth of the stream, copper concentrations increased and pH decreased with increased streamflow after the onset of winter rain and, unexpectedly, reached extreme values 1 or 2 months after peaks in the seasonal hydrographs. In contrast, aqueous zinc and sulfate concentrations were highest during low-flow periods. Spatial variation was assessed in 400 m of reach encompassing an acidic, metal-laden seep. At this seep, pH remained low (2-3) throughout the year, and copper concentrations were highest. In contrast, the zinc concentrations increased with downstream distance. These spatial patterns were caused by immobilization of copper by hydrous ferric oxides in benthic sediments, coupled with increasing downstream supply of zinc from groundwater seepage.

Seasonal and spatial patterns of metals at a restored copper mine site II. Copper in riparian soils and Bromus carinatus shoots.

Soil and plants were sampled throughout winter and spring near a perennial stream traversing a restored mine site in a winter-rainy climate. Within 1m of an acidic reach of the stream, soil had pH 3-5 and 50-100 microg/g "bioavailable" copper (extractable with 0.01 M CaCl2). Soil 2-3 m from the stream had pH 5-8 and lower (less than 3 microg/g) bioavailable copper. "Oxide-bound" copper (extractable with 2N HCl) was 50-100 microg/g at most locations. Copper concentrations in the shoots of field-collected Bromus carinatus declined from 20 microg/g in winter to 2 microg/g in spring at all sampling sites. A similar temporal pattern was found in plants grown under controlled conditions. Thus B. carinatus has a developmental program for control of shoot copper concentration, causing a seasonally-varying pattern of copper phytoaccumulation over a large range of copper availability in the soil.

Rates of root and organism growth, soil conditions, and temporal and spatial development of the rhizosphere.

BACKGROUND: Roots growing in soil encounter physical, chemical and biological environments that influence their rhizospheres and affect plant growth. Exudates from roots can stimulate or inhibit soil organisms that may release nutrients, infect the root, or modify plant growth via signals. These rhizosphere processes are poorly understood in field conditions. SCOPE AND AIMS: We characterize roots and their rhizospheres and rates of growth in units of distance and time so that interactions with soil organisms can be better understood in field conditions. We review: (1) distances between components of the soil, including dead roots remnant from previous plants, and the distances between new roots, their rhizospheres and soil components; (2) characteristic times (distance(2)/diffusivity) for solutes to travel distances between roots and responsive soil organisms; (3) rates of movement and growth of soil organisms; (4) rates of extension of roots, and how these relate to the rates of anatomical and biochemical ageing of root tissues and the development of the rhizosphere within the soil profile; and (5) numbers of micro-organisms in the rhizosphere and the dependence on the site of attachment to the growing tip. We consider temporal and spatial variation within the rhizosphere to understand the distribution of bacteria and fungi on roots in hard, unploughed soil, and the activities of organisms in the overlapping rhizospheres of living and dead roots clustered in gaps in most field soils. CONCLUSIONS: Rhizosphere distances, characteristic times for solute diffusion, and rates of root and organism growth must be considered to understand rhizosphere development. Many values used in our analysis were estimates. The paucity of reliable data underlines the rudimentary state of our knowledge of root-organism interactions in the field.

Moving with the flow: what transport laws reveal about cell division and expansion.

This material was presented as a keynote talk for the symposium, "Crosstalk between cell division and expansion," organized by G.T.S. Beemster and H. Tsukaya at the International Botanical Congress, Vienna in July, 2005. The review focuses on the utility of continuity equations to understand relationships among cell size, division and expansion; insights from Lagrangian or cell-specific descriptions of developmental variables; and a growth-diffusion equation to show effects of root growth zones on the surrounding soil.

Aphid infestation causes different changes in carbon and nitrogen allocation in alfalfa stems as well as different inhibitions of longitudinal and radial expansion.

Alfalfa (Medicago sativa) stem elongation is strongly reduced by a pea aphid (Acyrthosiphon pisum Harris) infestation. As pea aphid is a phloem feeder that does not transmit virus or toxins, assimilate withdrawal is generally considered as the main mechanism responsible for growth reduction. Using a kinematic analysis, we investigated the spatial distributions of relative elemental growth rates of control and infested alfalfa stems. The water, carbon, and nitrogen contents per unit stem length were measured along the growth zone. Deposition rates and growth-sustaining fluxes were estimated from these patterns. Severe short-term aphid infestation (200 young adults over a 24-h period) induced a strong and synchronized reduction in rates of elongation and of water and carbon deposition. Reduced nitrogen content and associated negative nitrogen deposition rates were observed in some parts of the infested stems, especially in the apex. This suggested a mobilization of nitrogen from the apical part of the growth zone, converted from a sink tissue into a source tissue by aphids. Calculation of radial growth rates suggested that aphid infestation led to a smaller reduction in radial expansion than in elongation. Together with earlier observations of long-lasting effects of a short-term infestation, this supports the hypothesis that in addition to nutrient withdrawal, a thigmomorphogenesis-like mechanism is involved in the effect of aphid infestation on stem growth.

The importance of frictional interactions in maintaining the stability of the twining habit.

The stability of twining vines under gravitational loads suggests an important role for friction. The coefficient of friction, µ, between vine stems and wood is high, often five times greater than between leather and wood, as determined by slip tests on an inclined plane. Stem trichomes function like ratchets to facilitate climbing upward (or to facilitate slipping if the stem is inverted). A mathematical model predicts large masses (kg) must be applied to the base of a twining vine to cause slipping. Vines slip as predicted when µ is low and arc length on the pole is short, and they break before slipping when µ is large or arc length is long. In contrast, twining vines are unstable in compression, collapsing when small masses (<10 g) are hung from the top of the vine. However, if the loads are applied below the uppermost gyre, the stabilizing tensional effect dominates. Therefore, in nature vines twining on a cylindrical support are stable under gravitational loads, unless these loads occur near the apex. A corollary is that a short apical coil can hold up large masses of maturing shoot.

Hydraulics of plant growth.

Multicellular plants rely on growth in localised regions that contain small, undifferentiated cells and may be many millimetres from the nearest differentiated xylem and phloem. Water and solutes must move to these small cells for their growth. Increasing evidence indicates that after exiting the xylem and phloem, water and solutes are driven to the growing cells by gradients in water potential and solute potential or concentration. The gradients are much steeper than in the vascular transport system and can change in magnitude or suffer local disruption with immediate consequences for growth. Their dynamics often obscure how turgor drives wall extension for growth, and different flow paths for roots and shoots have different dynamics. In this review, the origins of the gradients, their mode of action and their consequences are discussed, with emphasis on how their dynamics affect growth processes.