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An application test of the crop calendar portion of a corn (maize) stress indicator model developed by the early warning, crop condition assessment component of AgRISTARS was performed over the corn for grain producing regions of the U.S.S.R. during the 1980 crop year using real data. Performance of the crop calendar submodel was favorable; efficiency gains in meteorological data analysis time were on a magnitude of 85 to 90 percent.
The role of calcium redistribution in the responding region of the root is examined, however, the potential connection between calcium and auxin redistribution in the elongation zone is not found. The following items are examined: (1) the effect of gravity on calcium movement across the elongation zone; (2) the effect of gravity on auxin movement across the elongation zone; and (3) the effect of calcium on auxin movement across the elongation zone. It is indicated that gravistimulation induces a physiological asymmetry in the auxin transport system of maize roots and that calcium increases the total transport of auxin across the root. Gravistimulation is apparently necessary for the enhancing effect of calcium on lateral auxin movement, and it is possible that the preferential downward movement of calcium across the elongation zone of gravistimulated roots plays a role in establishing the auxin asymmetry proposed to cause positive gravitropic curvature.
Work of the Early Warning and Crop Condition Assessment (EW/CCA) project, one of eight projects in the Agriculture and Resources Inventory Surveys Through Aerospace Remote Sensing (AgRISTARS), is reviewed. Its mission, to develop and test remote sensing techniques that enhance operational methodologies for crop condition assessment, was in response to initiatives issued by the Secretary of Agriculture. Meteorologically driven crop stress indicator models have been developed or modified for wheat, maize, grain sorghum, and soybeans. These models provide early warning alerts of potential or actual crop stresses due to water deficits, adverse temperatures, and water excess that could delay planting or harvesting operations. Recommendations are given for future research involving vegetative index numbers and the NOAA and Landsat satellites.
The suitability of the Simple Biosphere (SiB) model of Sellers et al. (1986) for calculation of the surface fluxes for use within general circulation models is assessed. The structure of the SiB model is described, and its performance is evaluated in terms of its ability to realistically and accurately simulate biophysical processes over a number of test sites, including Ruthe (Germany), South Carolina (U.S.), and Central Wales (UK), for which point biophysical and micrometeorological data were available. The model produced simulations of the energy balances of barley, wheat, maize, and Norway Spruce sites over periods ranging from 1 to 40 days. Generally, it was found that the model reproduced time series of latent, sensible, and ground-heat fluxes and surface radiative temperature comparable with the available data.
A conventional radiometer is used to identify the hot spot (the peak in reflected radiation in the retrosolar direction) of vegetation. A multiwavelength-band radiometer collected radiances on fully grown dense wheat and maize canopies on several clear sunny days. It is noted that the hot spot is difficult to detect in the near IR wavelengths because the shadows are much darker. In general, the retrosolar brightness is found to be higher for smaller sun polar angles than for larger angles.
There is general agreement that, in roots, the primary detection of the gravitropic signal occurs in the columella cells of the cap and that this results in the generation of a signal that moves into the elongation zone causing the asymmetric growth that leads to downward curvature. Recent work has generated considerable evidence that indicates that auxin is the ultimate mediator of differential growth during root (and shoot) gravitropism. Our studies of the time course of curvature, auxin redistribution and/or adaptation, and electrical potential changes in maize roots have led to the following generalizations: (1) downward curvature begins 18 to 32 min following gravistimulation; (2) asymmetric auxin redistribution across the root cap begins at about the same time as curvature or perhaps slight earlier; (3) there is a lag of approx. 15 min in the response of roots to applied auxin; and (4) gravi-induced changes in intracellular potentials of cortical cells within the elongation zone occur within 30 s following stimulation.
In this experiment, layers of radiation detectors and biological specimens, bacterial spores (Bacillus subtillis), shrimp eggs (Altemia salina), and maize seeds (Zea mays) are sandwiched together in the Radiation Monitoring Container. The detectors, sheets of plastic materials, record the nuclear track of cosmic radiation. The dosimeter package contains conventional detectors made of materials such as lithium fluoride or magnesium-silica-terbium. The thermoluminescent materials (TLD) will, when moderately heated, emit luminescent photons linearly depending upon the dose of radiation received. The experiment, enclosed in a box-like container, is mounted on the aft end cone of the Spacelab, the area where the shielding is somewhat less than other locations.
The growth and development of more than 500 Zea mays seeds flown on Long Duration Exposure Facility (LDEF) were studied. Somatic mutations, including white-yellow stripes on leaves, dwarfing, change of leaf sheath color or seedling color were observed in plants developed from these seeds. When the frequency of white-yellow formation was used as the endpoint and compared with data from ground based studies, the dose to which maize seeds might be exposed during the flight was estimated to be equivalent to 635 cGy of gamma rays. Seeds from one particular holder gave a high mutation frequency and a wide mutation spectrum. White-yellow stripes on leaves were also found in some of the inbred progenies from plants displayed somatic mutation. Electron microscopy studies showed that the damage of chloroplast development in the white-yellow stripe on leaves was similar between seeds flown on LDEF and that irradiated by accelerated heavy ions on ground.
Environmental signals such as light and gravity control many aspects of plant growth and development. In higher plants, the directional growth of an organ in response to stimuli such as gravity and light is considered a tropic movement. Such movement could be either positive or negative with respect to a specific stimulus. In general, stems show a positive response to light and negative response to gravity. In contrast, most roots show a positive response to gravity and a negative response to light. Investigations on plant tropism date back a century when Darwin studied the phototropic responses of maize seedlings (Darwin). Although the precise mechanism of signal perception and transduction in roots is not understood, Darwin recognized over 100 years ago that the root cap is the probable site of signal perception. He discovered that the removal of the root cap eliminates the ability of roots to respond to gravity. Other investigators have since confirmed Darwin's observation (Konings; Evans et al.). In recent years, especially with the advent of the U.S. Space Program, there has been a renewed interest in understanding how plants respond to extracellular signals such as gravity (Halstead and Dutcher). Studies on the mechanisms involved in perception and transduction of gravity signal by roots would ultimately help us to better understand gravitropism and also to grow plants under microgravity conditions as in space. In this chapter, we restrict ourselves to the role of calcium in transduction of the gravity signal. In doing so, emphasis is given to the role of calcium-modulated proteins and their role in signal transduction in gravitropism. Detailed reviews on various other aspects of gravitropism (Scott, Torrey, Wilkins, Fim and Digby, Feldman, Pickard, Moore and Evans, Halstead and Dutcher, Poovaiah et al.) and on the role of calcium as a messenger in signal transduction in general have been published (Helper and Wayne, Poovaiah and Reddy, Roberts and Hartnon, Bowler and Chua, Gilroy and Trewavas). Plant roots have been widely used to study the transduction of gravity and light signals (Poovaiah et al., Roux and Serlin). Most roots show positive gravitropic response in either dark or light. However, roots of some varieties of plants (e.g., Zea mays L., cv Merit, and Zea rwvs L., cv Golden Cross Bantam 70) show positive gravitropic response only in light (Feldman, Miyazaki et al.). Investigations from various laboratories indicate that calcium acts as a messenger in transducing gravity and light signals in plant roots(Pickard, Evans et al., Pooviah et al.).
A model combining the rate of carbon assimilation with water and energy balance equations has been run using satellite and ancillary data for a period of 60 months (January 1986 to December 1990). Calculations for the Gediz basin area give mean annual evaporation as 395 mm, which is composed of 45% transpiration, 42% soil evaporation and 13% interception. The coefficient of interannual variation of evaporation is found to be 6%, while that for precipitation and net radiation are, respectively, 16% and 2%, illustrating that net radiation has an important effect in modulating interannual variation of evaporation. The mean annual water use efficiency (i.e., the ratio of net carbon accumulation and total evaporation) is ca. 1 g/sq m/mm, and has a coefficient of interannual variation of 5%. A comparison of the mean water use efficiency with field observations suggests that evaporation over the area is utilized well for biomass production. The reference crop evaporation for irrigated areas has annual mean and coefficient of variation as, respectively, 1176 mm and 3%. The total evaporation during three summer months of peak evaporation (June-August) is estimated to be about 575 mm for irrigated crops like maize and cotton. Seasonal variations of the fluxes are presented.
The plant hormone auxin affects cell elongation in both roots and shoots. In roots, the predominant action of auxin is to inhibit cell elongation while in shoots auxin, at normal physiological levels, stimulates elongation. The question of whether the primary receptor for auxin is the same in roots and shoots has not been resolved. In addition to its action on cell elongation in roots and shoots, auxin is transported in a polar fashion in both organs. Although auxin transport is well characterized in both roots and shoots, there is relatively little information on the connection, if any, between auxin transport and its action on elongation. In particular, it is not clear whether the protein mediating polar auxin movement is separate from the protein mediating auxin action on cell elongation or whether these two processes might be mediated by one and the same receptor. We examined the identity of the auxin growth receptor in roots and shoots by comparing the response of roots and shoots of the grass Zea mays L. and the legume Vigna mungo L. to indole-3-acetic acid, 2-naphthoxyacetic acid, 4,6-dichloroindoleacetic acid, and 4,7-dichloroindoleacetic acid. We also studied whether or not a single protein might mediate both auxin transport and auxin action by comparing the polar transport of indole-3-acetic acid and 2-naphthoxyacetic acid through segments from Vigna hypocotyls and maize coleoptiles. For all of the assays performed (root elongation, shoot elongation, and polar transport) the action and transport of the auxin derivatives was much greater in the dicots than in the grass species. The preservation of ligand specificity between roots and shoots and the parallels in ligand specificity between auxin transport and auxin action on growth are consistent with the hypothesis that the auxin receptor is the same in roots and shoots and that this protein may mediate auxin efflux as well as auxin action in both organ types.
Modulation of cytosolic calcium levels in both plants and animals is achieved by a system of Ca2+-transport and storage pathways that include Ca2+ buffering proteins in the lumen of intracellular compartments. To date, most research has focused on the role of transporters in regulating cytosolic calcium. We used a reverse genetics approach to modulate calcium stores in the lumen of the endoplasmic reticulum. Our goals were two-fold: to use the low affinity, high capacity Ca2+ binding characteristics of the C-domain of calreticulin to selectively increase Ca2+ storage in the endoplasmic reticulum, and to determine if those alterations affected plant physiological responses to stress. The C-domain of calreticulin is a highly acidic region that binds 20-50 moles of Ca2+ per mole of protein and has been shown to be the major site of Ca2+ storage within the endoplasmic reticulum of plant cells. A 377-bp fragment encoding the C-domain and ER retention signal from the maize calreticulin gene was fused to a gene for the green fluorescent protein and expressed in Arabidopsis under the control of a heat shock promoter. Following induction on normal medium, the C-domain transformants showed delayed loss of chlorophyll after transfer to calcium depleted medium when compared to seedlings transformed with green fluorescent protein alone. Total calcium measurements showed a 9-35% increase for induced C-domain transformants compared to controls. The data suggest that ectopic expression of the calreticulin C-domain increases Ca2+ stores, and that this Ca2+ reserve can be used by the plant in times of stress.
Characterization of gravitropic bending in the maize stem pulvinus, a tissue that functions specifically in gravity responses, demonstrates that the pulvinus is an ideal system for studying gravitropism. Gravistimulation during the second of three developmental phases of the pulvinus induces a gradient of cell elongation across the non-growing cells of the pulvinus, with the most elongation occurring on the lower side. This cell elongation is spatially and temporally separated from normal internodal cell elongation. The three characterized growth phases in the pulvinus correspond closely to a specialized developmental sequence in which structural features typical of cells not fully matured are retained while cell maturation occurs in surrounding internodal and nodal tissue. For example, the lignification of supporting tissue and rearrangement of transverse microtubules to oblique that occur in the internode when cell elongation ceases are delayed for up to 10 d in the adjacent cells of the pulvinus, and only occurs as a pulvinus loses its capacity to respond to gravistimulation. Gravistimulation does not modify this developmental sequence. Neither wall lignification nor rearrangement of transverse microtubules occurs in the rapidly elongating lower side or non-responsive upper side of the pulvinus until the pulvinus loses the capacity to bend further. Gravistimulation does, however, lead to the formation of putative pit fields within the expanding cells of the pulvinus.
The abscisic-acid (ABA) content of roots of the carotenoid-deficient w-3, vp-5, and vp-7 mutants of Z. mays was analyzed using gas chromatography-mass spectrometry with an analysis sensitivity of 6 ng ABA g-1 fresh weight (FW). Roots of normal seedlings of the same lines were characterized by the following amounts of ABA (as ng ABA g-1 FW, +/- standard deviation): w-3, 279 +/- 43; vp-5, 237 +/- 26; vp-7, 338 +/- 61. We did not detect any ABA in roots of any of the mutants. Thus, the lack of carotenoids in these mutants correlated positively with the apparent absence of ABA. Primary roots of normal and mutant seedlings were positively gravitropic, with no significant differences in the curvatures of roots of normal as compared with mutant seedlings. These results indicate that ABA 1) is synthesized in maize roots via the carotenoid pathway, and 2) is not necessary for positive gravitropism by primary roots of Z. mays.
Plants grown in microgravity are subject to many environmental stresses that may promote microbial growth and result in disease symptoms. Wheat (cv. Super Dwarf) recovered from an 8-day mission aboard a NASA (National Aeronautics and Space Administration) space shuttle showed disease symptoms, including girdling of leaf sheaths and chlorosis and necrosis of leaf and root tissues. A Neotyphodium species was isolated from the seed and leaf sheaths of symptomatic wheat used in the spaceflight mission. Certain isozymes of a peroxidase unique to extracts from the microgravity-grown plants were observed in extracts from earth-grown Neotyphodium-infected plants but were not present in noninfected wheat. The endophytic fungus was eliminated from the wheat seed by prolonged heat treatment at 50 degrees C followed by washes with water at 50 degrees C. Plants from wheat seed infected with the Neotyphodium endophyte were symptomless when grown under greenhouse conditions, whereas symptoms appeared after only 4 days of growth in closed containers. Disease spread from an infected plant to noninfected plants in closed containers. Dispersion via spores was found on asymptomatic plants at distances of 7 to 18 cm from infected plants. The size and shape of the conidia, mycelia, and phialide-bearing structures and the ability to grow rapidly on carbohydrates, especially xylose, resembled the characteristics of N. chilense, which is pathogenic on orchard grass, Doctylis glomerati. The Neotyphodium wheat isolate caused disease symptoms on other cereals (wheat cv. Malcolm, orchard grass, barley, and maize) grown in closed containers.
The maize Ac-Ds transposable element family has been used as the basis of transposon mutagenesis systems that function in a variety of plants, including Arabidopsis. We have developed modified transposons and methods which simplify the detection, cloning and analysis of insertion mutations. We have identified and are analyzing two plant lines in which genes expressed either in the root cap cells or in the quiescent cells, cortex/endodermal initial cells and columella cells of the root cap have been tagged with a transposon carrying a reporter gene. A gene expressed in root cap cells tagged with an enhancer-trap Ds was isolated and its corresponding EST cDNA was identified. Nucleotide and deduced amino acid sequences of the gene show no significant similarity to other genes in the database. Genetic ablation experiments have been done by fusing a root cap-specific promoter to the diphtheria toxin A-chain gene and introducing the fusion construct into Arabidopsis plants. We find that in addition to eliminating gravitropism, root cap ablation inhibits elongation of roots by lowering root meristematic activities.
Previous work has shown that microtubule (MT) reorientation follows the onset of growth inhibition on the lower side of graviresponding roots, indicating that growth reduction can occur independently of MT reorientation. To test this observation further, we examined whether the reduction in growth in response to osmotic stress is correlated with MT reorientation. The distribution and rate of growth in maize roots exposed to 350 mOsm sorbitol and KCl or 5 mM Mes/Tris buffer were measured with a digitizer. After various times roots were processed for indirect immunofluorescence microscopy. Application of sorbitol or KCl had no effect on the organization of MTs in the apical 2 mm of the root but resulted in striking and different effects in the basal region of the root. Sorbitol treatment caused rapid appearance of oval to circular holes in the microtubular array that persisted for at least 9 h. Between 30 min and 4 h of submersion in KCl, MTs in cortical cells 4 mm and farther from the quiescent center began to reorient oblique to the longitudinal axis. After 9 h, the alignment of MTs had shifted to parallel to the root axis but MTs of the epidermal cells remained transverse. In KCl-treated roots MT reorientation appeared to follow a pattern of development similar to that in controls but without elongation. Our data provide additional evidence that MT reorientation is not the cause but a consequence of growth inhibition.
Two key issues in studies of the nature of the gravitropic response in roots have been the determination of the precise pattern of differential elongation responsible for downward bending and the identification of the cells that show the initial motor response. The main approach for examining patterns of differential growth during root gravitropic curvature has been to apply markers to the root surface and photograph the root at regular intervals during gravitropic curvature. Although these studies have provided valuable information on the characteristics of the gravitropic motor response in roots, their labor intensive nature limits sample size and discourages both high frequency of sampling and depth of analysis of surface expansion data. In this brief review we describe the development of computer-based video analysis systems for automated measurement of root growth and shape change and discuss some key features of the root gravitropic response that have been revealed using this methodology. We summarize the capabilities of several new pieces of software designed to measure growth and shape changes in graviresponding roots and describe recent progress in developing analysis systems for studying the small, but experimentally popular, primary roots of Arabidopsis. A key finding revealed by such studies is that the initial gravitropic response of roots of maize and Arabidopsis occurs in the distal elongation zone (DEZ) near the root apical meristem, not in the main elongation zone. Another finding is that the initiation of rapid elongation in the DEZ following gravistimulation appears to be related to rapid membrane potential changes in this region of the root. These observations have provided the incentive for ongoing studies examining possible links between potential growth modifying factors (auxin, calcium, protons) and gravistimulated changes in membrane potential and growth patterns in the DEZ.