Rodolfo A. Sánchez

ENVIRONMENTAL CONTROL OF DORMANCY IN WEED SEED BANKS IN SOIL

Dormancy is a common attribute of many weed seed populations and this usually hampers the task of predicting timing and extent of emergence of weeds. Both the number of established plants and the timing of emergence of a weed are strongly related to the dynamics of dormancy release of the seed population. In this paper, we discuss the different factors that affect dormancy in weed seed banks in soil, aiming to set a conceptual basis that will facilitate the construction of predictive models. From the long list of factors that are known to control dormancy under field conditions, we distinguish those that modify the dormancy level of the population (i.e. soil temperature and soil hydric conditions) from those that terminate dormancy or in other words, remove the ultimate constraints for seed germination once the degree of dormancy is sufficiently low (i.e. light, fluctuating temperatures, nitrate concentration). We also discuss the effect of agricultural practices on dormancy of weed seed populations, making reference to studies that have evinced clearly the factor(s) involved in determining a particular pattern of response. Overall, we stress the importance of clarifying, both qualitatively and quantitatively, the interaction between soil thermal and hydric conditions in the modification of the degree of dormancy of seed populations. Similarly, it is essential that we understand the extent to which such changes in dormancy comprise changes in sensitivity to factors that terminate dormancy. # 2000 Elsevier Science B.V. All rights reserved.

Tillering responses to enrichment of red light beneath the canopy in a humid natural grassland

(1) Enrichment of red light at the base of dallisgrass (Paspalum dilatatum Poir.) and smutgrass (Sporobolus indices [L.] R.Br.) plants in a dense humid natural grassland was obtained by use of small light-emitting diodes around the crown of each grass plant. (2) Results of this study supported the hypothesis that modification in light quality by a dense grassland canopy was detrimental to tiller populations. Tillering rates were increased and tiller death delayed until the end of the growing season when additional red light was supplied to the crown of plants. (3) The photocontrol of axillary bud activity may be part of a system whereby the tiller dynamics would be related to resources availability. (4) These results also suggest that maintenance of a grass sward at a high leaf area index might not be a good management strategy for long-term productivity.

Phytochrome A Mediates the Promotion of Seed Germination by Very Low Fluences of Light and Canopy Shade Light in Arabidopsis.

Seeds of the wild type (WT) and of the phyA and phyB mutants of Arabidopsis thaliana were exposed to single red light (R)/far-red light (FR) pulses predicted to establish a series of calculated phytochrome photoequilibria (Pfr/P). WT and phyB seeds showed biphasic responses to Pfr/P. The first phase, i.e. the very-low-fluence response (VLFR), occurred below Pfr/P = 10-1%. The second phase, i.e. the low-fluence response, occurred above Pfr/P = 3%. The VLFR was similarly induced by either a FR pulse saturating photoconversion or a subsaturating R pulse predicted to establish the same Pfr/P. The VLFR was absent in phyA seeds, which showed a strong low-fluence response. In the field, even brief exposures to the very low fluences of canopy shade light (R/FR ratio < 0.05) promoted germination above dark controls in WT and phyB seeds but not in the phyA mutant. Seeds of the phyA mutant germinated normally under canopies providing higher R/FR ratios or under deep canopy shade light supplemented with R from light-emitting diodes. We propose that phytochrome A mediates VLFR of A. thaliana seeds.

Phytochromes and seed germination

The control of seed germination by red and far-red light is one of the earliest documented phytochrome-mediated processes Phytochrome is now known to be a small family of photoreceptors whose apoproteins are encoded by different genes Phytochrome B (phyB) is present in dry seeds and affects germination of dark imbibed seeds but other phytochromes could also be involved Phytochrome A (phyA) appears after several hours of imbibition and mediates very-low-fluence responses PhyB and other phytochromes different from phyA mediate the classical low-fluence responses The phytochrome involved in high-irradiance responses of seed germination (inhibition of germination under continuous far-red) has not been unequivocally established, although phyA is the most likely candidate Phytochrome can affect embryo growth capacity and/or the constraint imposed by the tissues surrounding the embryo At least in some species, gibberellins participate in the signalling process In the field, phyA has been implicated in the perception of light during soil cultivations, and phyB would be involved in the perception of red/far-red ratios associated with the presence of gaps in the canopy This review describes recent advances in phytochrome research, particularly those derived from the analysis of germination in specific mutants, and their connection with traditional observations on phytochrome control of seed germination

Superoxide anion and hydrogen peroxide metabolism in soybean embryonic axes during germination

The total rate of mitochondrial O2- production in the presence of NADH as substrate increased from 200 to 1340 pmol/min per axis between 2 and 30 h of imbibition. The activities of the enzymes involved in hydroperoxide metabolism, e.g., superoxide dismutase, catalase, peroxidase and glutathione and ascorbate peroxidases, markedly changed during the germination of soybean embryonic axes. Superoxide dismutase was the enzymatic activity affected the most during the initial stages of germination. Intracellular O2- steady-state concentration, calculated from the rate of O2- production and superoxide dismutase activity, showed a 2-fold increase from 2 x 10(-8) M to 4 x 10(-8) M in germination phase I, declined in phase II to 2 x 10(-8) M and remained constant over the rest of the incubation period. The reaction of H2O2 and luminol catalyzed by Co2+ was utilized to measure H2O2 diffused out of the soybean axes after 5 to 10 min of incubation. The catalase-sensitive luminol emission of diffusates prepared from axes previously imbibed from 2 to 30 h corresponded to a H2O2 intracellular steady-state concentration in the range of 0.3 to 0.9 microM. The activity of metal-containing antioxidant enzymes was determined in the extracellular fluid. Cell wall peroxidase activity increased from 10 to 300 mumol/min per mg protein and appears as a potentially important pathway for H2O2 utilization. Hydrogen peroxide metabolism in soybean embryonic axes during early inhibition appears to have the following main features: (a) mitochondrial membranes are the most important source of cytosolic O2- and H2O2; (b) H2O2 is regulated at a steady-state concentration of 0.3-0.9 microM; (c) catalase is the main enzyme in terms of H2O2 utilization; (d) H2O2 exo-diffusion is quantitatively important destiny of intracellular H2O2; and (e) extracellular peroxidase located at the cell wall affords an enzymatic system able to use diffused H2O2.

The effect of plant density on tillering: The involvement of R/FR ratio and the proportion of radiation intercepted per plant

Abstract Plants of Paspalum dilatatum and Lolium multiflorum were grown in pots at different densities. Half of the plants at each density received additional red light at their bases by means of light-emitting diodes. Density reduced both the proportion of incident radiation intercepted per plant and the red/far-red ratio of the light at plant bases. Tillering decreased with density in both species. P. dilatatum plants also had fewer reproductive tillers and expanded leaves and a higher mortality of young vegetative tillers in denser stands. In both species the enrichment with red light did not increase tillering of isolated plants, but increased it in stands grown at relatively low densities with small degrees of mutual shading. At these densities P. dilatatum plants achieved the number of tillers of isolated plants. Moreover, red light increased vegetative tiller death in the P. dilatatum dense canopy. These results suggest that low red/far-red ratios might preclude morphogenic responses to density before an important depletion in energy availability takes place.

Increased Phytochrome B Alleviates Density Effects on Tuber Yield of Field Potato Crops

The possibility that reduced photomorphogenic responses could increase field crop yield has been suggested often, but experimental support is still lacking. Here, we report that ectopic expression of the Arabidopsis PHYB (phytochrome B) gene, a photoreceptor involved in detecting red to far-red light ratio associated with plant density, can increase tuber yield in field-grown transgenic potato (Solanum tuberosum) crops. Surprisingly, this effect was larger at very high densities, despite the intense reduction in the red to far-red light ratios and the concomitant narrowed differences in active phytochrome B levels between wild type and transgenics at these densities. Increased PHYB expression not only altered the ability of plants to respond to light signals, but they also modified the light environment itself. This combination resulted in larger effects of enhanced PHYB expression on tuber number and crop photosynthesis at high planting densities. The PHYB transgenics showed higher maximum photosynthesis in leaves of all strata of the canopy, and this effect was largely due to increased leaf stomatal conductance. We propose that enhanced PHYB expression could be used in breeding programs to shift optimum planting densities to higher levels.

Hormonal Regulation of Dormancy in Developing Sorghum Seeds

The role of abscisic acid (ABA) and gibberellic acid (GA) in determining the dormancy level of developing sorghum (Sorghum bicolor [L.] Moench.) seeds from varieties presenting contrasting preharvest sprouting behavior (Redland B2, susceptible; IS 9530, resistant) was investigated. Panicles from both varieties were sprayed soon after pollination with fluridone or paclobutrazol to inhibit ABA and GA synthesis, respectively. Fluridone application to the panicles increased germinability of Redland B2 immature caryopses, whereas early treatment with paclobutrazol completely inhibited germination of this variety during most of the developmental period. Incubating caryopses in the presence of 100 [mu]M GA4+7 overcame the inhibitory effect of paclobutrazol, but also stimulated germination of seeds from other treatments. IS 9530 caryopses presented germination indices close to zero until physiological maturity (44 d after pollination) in control and paclobutrazol-treated particles. However, fluridone-treated caryopses were released from dormancy earlier than control and paclobutrazol-treated caryopses. Incubation in the presence of GA4+7 stimulated germination of caryopses from all treatments. Our results support the proposition that a low dormancy level (which is related to a high preharvest sprouting susceptibility) is determined not only by a low embryonic sensitivity to ABA, but also by a high GA content or sensitivity.

Photomorphogenic processes in the agricultural environment

We review recent advances in our understanding of how photomorphogenic mechanisms influence processes of significance for agriculture, including weed seed germination, acclimation to solar UV‐B radiation, and plant interactions in canopies.

Morphological responses of Datura ferox L. seedlings to the presence of neighbours Their relationships with canopy microclimate

SummaryWe studied the effects of density on the dynamics of seedling growth and canopy microclimate within experimental stands composed of Datura ferox L. seedlings grown in individual pots. Interception of photosynthetically active radiation (PAR) by seedlings was evaluated either indirectly, by measuring leaf area, proportion of leaf area shaded by neighbouring individuals and laminar orientation with respect to sunlight, or directly, by measuring PAR at individual leaves at their natural angle of display. An integrating cylinder, with a geometry approximating that of a stem, was used within the canopies to measure the red:far-red (R:FR) ratio of the light flux from all compass points parallel to the soil surface. Seedlings responded rapidly (i.e. 1–2 weeks) to increased density by producing longer internodes and partitioning more dry matter to stems relative to leaves. These responses were observed before either PAR interception of growth rate were reduced by the presence of neighbours. Conversely, morphogenetic adjustment was preceded by a consistent effect of plant density on the R:FR ratio of the light received by the integrating cylinder. Air and soil temperature were not affected by density in these experiments. Differences in wind velocity within the canopy associated with plant density were avoided by the experimental procedure. The results support the idea that the drop in R:FR ratio of the light flux parallel to the ground — e.g. reflected sunlight — is an early signal that allows rapid adjustment of plant form to changes in canopy structure.