Differential aluminum resistance in Brachiaria species

Brachiaria are increasingly cropped in the tropics because these species combine good fodder quality and yield with high resistance to aluminum (Al) toxicity, an important stress factor in acidic soils. The mechanisms for the extraordinarily high resistance to Al toxicity in Brachiaria decumbens remain unclear. It has been suggested that the presence of a multiseriate exodermis might contribute to efficient Al exclusion in B. decumbens. However, no data concerning the root structure of less Al-resistant Brachiaria species have been reported. The aim of the present study was determine whether the exodermis is a distinctive feature of Al hyper-resistant B. decumbens compared with Brachiaria species with lower Al resistance. B. decumbens, B. brizantha, and B. ruziziensis were grown in nutrient solution without (control) or with 200 μM Al (32 μM Al³⁺ activity) for 96 h. Differences in the Al resistance were assessed using various indicators: Al-induced inhibition of root elongation, membrane damage, and the maintenance of nutrient homeostasis. Transversal root sections were examined using fluorescence microscopy to reveal the presence of an exodermis through auto-fluorescence. Aluminum resistance decreased in the order B. decumbens > B. brizantha > B. ruziziensis. Both the hyper-resistant B. decumbens and the moderately resistant B. brizantha were more efficient in Al-exclusion than the sensitive B. ruziziensis. Apoplastic barriers, in the form of a multiseriate exodermis, were constitutively present in B. decumbens, but not in Al-sensitive B. ruziziensis. Under control conditions, B. brizantha exhibited slightly auto-fluorescent epidermal cell walls, while under Al exposure auto-fluorescent deposits were observed in the intercellular spaces between the epidermal and sub-epidermal cell layers. The results provide circumstantial evidence of a role for apoplastic barriers in the Al resistance of B. decumbens and, to a lesser extent, in B. brizantha. Nonetheless, additional research is required to determine a causal relationship between the exodermal barrier and Al resistance.     

Aluminum tolerance measured by root growth and mucilage protection in Urochloa brizantha and Urochloa decumbens

Aluminum toxicity on root systems was analyzed through comparing root growth and evaluating the protective function of mucilage in Urochloa decumbens and Urochloa brizantha. Seedlings were grown in a solution with different concentrations of AlCl₃ and with mucilage removed or present. The root elongation rate, total length, number of roots and presence of aluminum at the root apex were measured. Root development was inhibited by aluminum and the elongation rate was maintained without any difference between the two species. A significant reduction in root length was found in U. brizantha. Aluminum did not influence root branching in either species and accumulated mostly in the rhizosphere of U. brizantha, where the mucilage has less of a protective function. The greatest aluminum tolerance was found in U. decumbens, observed through maintenance of total root system growth from lower to higher aluminum toxicity.     

Responses to drought of five Brachiaria species. I. Biomass production,leaf growth,root distribution,water use and forage quality

The introduction of African grasses in Neotropical savannas has been a key factor to improve pasture productivity. We compared the response of five Brachiaria species to controlled drought (DT) in terms of biomass yield and allocation, pattern of root distribution, plant water use, leaf growth, nutrient concentration and dry matter digestibility. The perennial C₄ forage grasses studied were B. brizantha (CIAT 6780), B. decumbens (CIAT 606), B. dictyoneura (CIAT 6133), B. humidicola (CIAT 679) and B. mutica. Two DT periods, which mimic short dry spells frequent in the rainy season, were imposed by suspending irrigation until wilting symptoms appeared. They appeared after 14 days in B. brizantha, B. decumbens and B. mutica, and after 28 days in B. humidicola and B. dictyoneura. The impossed drought stress was mild and only the largest grass, B. brizantha, showed reduced (23%) plant yield. The other grasses were able to adjust growth and biomass allocation in response to DT leaving total plant yield relatively unaffected. Brachiaria mutica, had a homogeneous root distribution throughout the soil profile. In the other species more than 80% of root biomass was allocated within the first 30 cm of the soil profile. Brachiaria brizantha and B. decumbens had the lowest proportion of roots below 50 cm. Drought caused a general reduction in root biomass. The shoot:root ratio in B. mutica and B. humidicola increased in response to DT at the expense of a reduction in root yield down to 50 cm depth. Although the total water volume utilized under DT was similar among grasses, the rate of water use was highest (0.25 l day^–1) in B. brizantha, B. decumbens and B. mutica and lowest (0.13 l day^–1) in B. humidicola and B. dictyoneura. In all species leaf expansion was reduced by DT but it was rapidly reassumed after rewatering. Drought increased specific leaf mass (SLM) only in B. brizantha compensating for leaf area reduction, but leaf area ratio (LAR) was unaffected in all species. In almost all grasses DT increased leaf N and K concentration and in vitro dry matter digestibility. The results indicate that B. brizantha, B. decumbens and to a lesser extent, B. mutica are better adapted to short dry periods, whereas B. humidicola and B. dictyoneura are better adapted to longer dry periods.     

Chromosomal distribution of H3K4me2, H3K9me2 and 5-methylcytosine: variations associated with polyploidy and hybridization in <Emphasis Type="Italic">Brachiaria</Emphasis> (Poaceae)

Key message

Assessment of chromosomal distribution of modified histones and 5-methylcytosine shown that there are diversification of chromosomal types among species of Brachiaria and its interspecific hybrids.

Abstract

Histone post-translational modifications and DNA methylation are epigenetic processes that are involved in structural and functional organization of the genome. This study compared the chromosomal distribution of modified histones and 5-methylcytosine (5-mCyt) in species and interspecific hybrids of Brachiaria with different ploidy levels and reproduction modes. The relation between H3K9me2 and 5-mCyt was observed in the nucleolus organizer region, centromeric central domain and pericentromeric region. H3K4me2 was detected in euchromatic domains, mainly in the terminal chromosomal regions. Comparison of chromosomal distribution among species and hybrids showed greater variation of chromosomal types for the H3K9me2 in B. decumbens (tetraploid and apomictic species) and the 963 hybrid, while, for the H3K4me2, the variation was higher in B. brizantha and B. decumbens (tetraploid and apomictic species) and 963 hybrid. The chromosome distribution of 5-mCyt was similar between B. brizantha and B. decumbens, which differ from the distribution observed in B. ruziziensis (diploid and sexual species). Significant alterations in DNA methylation were observed in the artificially tetraploidized B. ruziziensis and in the interspecific hybrids, possibly as result of hybridization and polyploidization processes. The monitoring of histone modifications and DNA methylation allowed categorizing nuclear and chromosomal distribution of these epigenetic marks, thus contributing to the knowledge of composition and structure of the genome/epigenome of Brachiaria species and hybrids. These data can be useful for speciation and genome evolution studies in genus Brachiaria, and represent important markers to explore relationships between genomes.

     

A comparative study on plant growth and root plasticity responses of two Brachiaria forage grasses grown in nutrient solution at low and high phosphorus supply

Brachiaria forage grasses are widely used for livestock production in the tropics. Signalgrass (Brachiaria decumbens cv. Basilisk, CIAT 606) is better adapted to low phosphorus (P) soils than ruzigrass (B. ruziziensis cv. Kennedy, CIAT 654), but the physiological basis of differences in low-P adaptation is unknown. We characterized morphological and physiological responses of signalgrass and ruzigrass to low P supply by growing both grasses for 30 days in nutrient solution with two levels of P supply using the hydroxyapatite pouch system. Ruzigrass produced more biomass at both levels of P supply whilst signalgrass appears to be a slower-growing grass. Both grasses increased biomass allocation to roots and had higher root acid phosphatase and phytase activities at low P supply. At low P supply, ruzigrass showed greater morphological plasticity as its leaf mass density and lateral root fraction increased. For signalgrass, morphological traits that are not responsive to variation in P supply might confer long-term ecological advantages contributing to its superior field persistence: greater shoot tissue mass density (dry matter content) might lower nutrient requirements while maintenance of lateral root growth might be important for nutrient acquisition in patchy soils. Physiological plasticity in nutrient partitioning between root classes was also evident for signalgrass as main roots had higher nutrient concentrations at high P supply. Our results highlight the importance of analyzing morphological and physiological trait profiles and determining the role of phenotypic plasticity to characterize differences in low-P adaptation between Brachiaria genotypes.     

Arbuscular Mycorrhizal Fungi and Arsenate Uptake by Brachiaria Grass (Brachiaria decumbens)

ABSTRACT Using a compartmentalized treatment technique, the role of arbuscular mycorrhizal fungi (AMF; Acaulospora scrobiculata) on arsenic (As) uptake and translocation in Brachiaria decumbens. Treatments consisted of a factorial arrangement of three As doses (0, 50, and 100 mg kg^?1) and the presence/absence of AMF inoculates. In the absence of AMF, B. decumbens did not show As accumulation, indicating the probable presence of tolerance mechanism via As exclusion by the roots. B. decumbens plants showed high AMF colonization levels, especially in the arsenic treatments, with AMF improving shoot and root growth independent of As concentrations. Arsenic accumulation occurred only with AMF inoculation. Phosphorous uptake was reduced in B. decumbens roots in the presence of arsenic with and without inoculation of AMF. Results suggest that B. decumbens can be used in phytoremediation procedures when inoculated with A. scrobiculata, although pasture formation should be strictly avoided in contaminated sites.     

Host plant resistance in Brachiaria grasses to the spittlebug Zulia colombiana

Twelve forage grass accessions including 11 accessions of Brachiaria Griseb, were evaluated in a glasshouse for host plant resistance to nymphs and tolerance to feeding damage caused by adults of Zulia colombiana (Lallemand) (Homoptera: Cercopidae). Resistance to nymphs was evaluated with a technique that provided uniform environmental conditions and abundant feeding sites. B. brizantha Stapf (cv. Marandú) was the most resistant of the accessions tested based on nymphal mortality, duration of nymphal stadia, and weight of adult females. Andropogon gayanus Kunth, resistant to spittlebug attack in the field, was susceptible under the conditions of this study. While growth habit and rooting characteristics may contribute to field resistance, other resistance factors are present within the genus Brachiaria, particularly in the case of B. brizantha cv. Marandú. The number of insect-days causing severe damage in the most tolerant species (B. dictyoneura Stapf and B. humidicola Schweick) was approximately six times greater than that necessary to cause the same level of damage to the most susceptible species (B. ruziziensis Germain & Evrard and B. decumbens Stapf). No difference was found in regrowth capacity between infested and noninfested plants within accessions. There was a significant positive correlation between number of insect-days causing severe damage (tolerance) and regrowth of infested plants.

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Résumé Brachiaria est une graminée fourragère prometteuse pour les sols tropicaux acides, saturés d'aluminium. Z. colombiana est un Cercopidae très répandu, limitant l'utilisation de Brachiaria en Amérique Latine. La résistance (antibiose et tolérance) à Z. colombiana, de Brachiaria d'origines diverses a été examinée. B. brizantha cv. Marandù s'est révélé le plus résistant d'après la forte mortalité larvaire, la prolongation du développement larvaire, et le poids réduit des femelles adultes de Z. colombiana. Andropogon gayanus, résistant dans la nature, s'est révélé sensible. Ces résultats suggèrent que cette résistance de A. gayanus dans la nature pourrait être due à la structure du végétal et à son mode de croissance. Dans le cas de B. brizantha cv. Marandù, des facteurs supplémentaires de résistance, mis en évidence à partir de différents modes de croissance, ont été éliminés, de façon à identifier les mécanismes de l'antibiose présents chez Brachiaria. Une grande gamme de résistance aux attaques alimentaires a été observée chez Brachiaria. Les plus résistants ont besoin de 6 fois plus de jours d'attaque par Z. colombiana pour provoquer les dégâts observés sur individus sensibles.

     

GISH-based comparative genomic analysis in Urochloa P. Beauv.

The genus Urochloa P. Beauv. [syn. Brachiaria (Trin.) Griseb.] comprises species of great economic relevance as forages. The genomic constitution for the allotetraploid species Urochloa brizantha (cv. Marandu) and Urochloa decumbens (cv. Basilisk) and the diploid Urochloa ruziziensis was previously proposed as BBB¹B¹, B¹B¹B²B² and B²B², respectively. Evidence indicates U. ruziziensis as the ancestral donor of genome B² in U. decumbens allotetraploidy, but the origin of the genomes B and B¹ is still unknown. There are diploid genotypes of U. brizantha and U. decumbens that may be potential ancestors of the tetraploids. The aim of this study was to determine the genomic constitution and relationships between genotypes of U. brizantha (2x and 4x), U. decumbens (2x and 4x) and U. ruziziensis (2x) via genomic in situ hybridization (GISH). Additionally, chromosome number and genome size were verified for the diploid genotypes. The diploids U. brizantha and U. decumbens presented 2n?=?2x?=?18 chromosomes and DNA content of 1.79 and 1.44 pg, respectively. The GISH analysis revealed high homology between the diploids U. brizantha and U. decumbens, which suggests relatively short divergence time. The GISH using genomic probes from the diploid accessions on the tetraploid accessions’ chromosomes presented similar patterns, highlighting the genome B¹ present in both of the tetraploids. Based on GISH results, the genomic constitution was proposed for the diploid genotypes of U. brizantha (B¹B¹) and U. decumbens (B¹′B¹′) and both were pointed as donors of genome B¹ (or B¹′), present in the allotetraploid genotypes.

     

Physical mapping of rDNA genes corroborates allopolyploid origin in apomictic Brachiaria brizantha

Brachiaria (Trin.) Griseb belongs to the family Poaceae, and within the genus, apomixis or sexuality is present in different accessions of the same species. The majority of Brachiaria species are polyploid and apomictic, making strategies for crop improvement by breeding very intricate. In spite of the high frequency of apomictic polyploids, the relationship of polyploidy and hybridization with apomixis in Brachiaria is still unclear. Further analysis requires detailed knowledge regarding the genomic composition of the polyploids. The present work introduces the use of fluorescent in situ hybridization (FISH) into cytogenetic analysis of Brachiaria. Physical mapping of heterologous rDNA sequences, associated with conventional karyotyping of the B. brizantha diploid sexual (BRA 002747) and the tetraploid apomictic (BRA000591) accessions, provided evidence of the latter being of allopolyploid origin. Based on our results and on previous knowledge on apomixis in B. brizantha, we suggest that the origin of apomixis was probably a consequence of hybridization.     

Influence of inoculum density and aeration volume on biomass and bioactive compound production in bulb-type bubble bioreactor cultures of Eleutherococcus koreanum Nakai

This study deals with the effects of initial inoculum density and aeration volume on biomass and bioactive compound production in adventitious roots of Eleutherococcus koreanum Nakai in bulb-type bubble bioreactors (3-L capacity). While the fresh and dry weights of the roots increased with increasing inoculum density, the highest percentage dry weight and accumulation of total target compounds (eleutheroside B and E, chlorogenic acid, total phenolics, and flavonoids) were noted at an inoculum density of 5.0 g L⁻¹. Poor aeration volume (0.05 vvm) stunted root growth, and high aeration volume (0.4 vvm) caused physiological disorders. Moreover, an inoculum density of 5.0 g L⁻¹ and an aeration volume of 0.1 vvm resulted in the highest concentration of total target compounds and least root death. Such optimization of culture conditions will be beneficial for the large-scale production of E. koreanum biomass and bioactive compounds.     

Aluminium-induced changes in the profiles of both organic acids and phenolic substances underlie Al tolerance in Rumex acetosa L.

The common sorrel, Rumex acetosa L. is well adapted to acid mineral soils with high availability of phytotoxic Al species. The mechanisms of Al resistance in this species are not established. Our goal was to assess the possible implications of organic acids and phenolic substances in Al detoxification in roots and shoots of this plant. R. acetosa plants were exposed in nutrient solution (pH 4.3) to a non-growth reducing Al concentration of 50 μM Al for 5 days. Exclusion of Al from root tips was visualized by haematoxylin staining. Tissue Al and Ca concentrations were analysed by ICP ES. Root and shoot concentrations of organic acids and phenolic substances were analysed by HPLC. A time-dependent (model II type) Al exclusion pattern in root tips was observed. Nonetheless, high Al concentrations accumulated in roots (1170 μg/g) and shoots (275 μg/g). Aluminium supply enhanced root citrate concentrations but decreased shoot organic acid levels. Aluminium induced high levels of anthraquinone in roots and of catechol, catechin and rutin in shoots. Aluminium resistance in R. acetosa implies both exclusion of Al from root tips and tolerance to high Al tissue concentrations. Citrate in roots and phenolics in shoots may bind Al in non-toxic form. Anthraquinones, as strong antioxidants, may play a role in a general defence response to the root stress.     

Phosphorus enhances Al resistance in Al-resistant Lespedeza bicolor but not in Al-sensitive L. cuneata under relatively high Al stress

Background and Aims

Aluminium (Al) toxicity and phosphorus (P) deficiency often co-exist in acidic soils and limit crop production worldwide. Lespedeza bicolor is a leguminous forage species that grows very well in infertile, acidic soils. The objective of this study was to investigate the effects of Al and P interactions on growth of Lespedeza and the distributions of Al and P in two different Al-resistant species, and to explore whether P can ameliorate the toxic effect of Al in the two species.

Methods

Two species, Lespedeza bicolor and L. cuneata, were grown for 30 d with alternate Al and P treatments in a hydroponics system. Harvested roots were examined using a root-system scanner, and the contents of Al, P and other nutrient elements in the plants were determined using inductively coupled plasma-atomic emission spectroscopy (ICP-AES). Haematoxylin staining was used to observe the distribution of Al in the roots of seedlings. After pre-culture with or without P application, organic acids in the exudates of roots exposed to Al were held in an anion-exchange resin, eluted with 2 m HCl and then analysed using high-performance liquid chromatography (HPLC).

Key Results

Lespedeza bicolor exhibited a stronger Al resistance than did L. cuneata; Al exclusion mechanisms may mainly be responsible for resistance. P application alleviated the toxic effect of Al on root growth in L. bicolor, while no obvious effects were observed in L. cuneata. Much less Al was accumulated in roots of L. bicolor than in L. cuneata after P application, and the P contents in both roots and shoots increased much more for L. bicolor than for L. cuneata. Lespedeza bicolor showed a higher P/Al ratio in roots and shoots than did L. cuneata. P application decreased the Al accumulation in root tips of L. bicolor but not in L. cuneata. The amount of Al-induced organic acid (citrate and malate) exudation from roots pre-cultured with P was much less than from roots without P application; no malate and citrate exudation was detected in L. cuneata.

Conclusions

P enhanced Al resistance in the Al-resistant L. bicolor species but not in the Al-sensitive L. cuneata under relatively high Al stress, although P in L. cuneata might also possess an alleviative potential. Enhancement of Al resistance by P in the resistant species might be associated with its more efficient P accumulation and translocation to shoots and greater Al exclusion from root tips after P application, but not with an increased exudation of organic acids from roots.Key words: Lespedeza bicolor, L. cuneata, Al toxicity, Al resistance, root morphology, phosphorus     

Location of low copy genes in chromosomes of <Emphasis Type="Italic">Brachiaria</Emphasis> spp.

Repetitive DNA sequences have been widely used in cytogenetic analyses. The use of gene sequences with a low-copy-number, however, is little explored especially in plants. To date, the karyotype details in Brachiaria spp. are limited to the location of rDNA sites. The challenge lies in developing new probes based on incomplete sequencing data for the genus or complete sequencing of related species, since there are no model species with a sequenced genome in Brachiaria spp. The present study aimed at the physical location of conserved genes in chromosomes of Brachiaria ruziziensis, Brachiaria brizantha, and Brachiaria decumbens using RNAseq data, as well as sequences of Setaria italica and Sorghum bicolor through the fluorescent in situ hybridization technique. Five out of approximately 90 selected sequences generated clusters in the chromosomes of the species of Brachiaria studied. We identified genes in synteny with 5S and 45S rDNA sites, which contributed to the identification of chromosome pairs carrying these genes. In some cases, the species of Brachiaria evaluated had syntenic segments conserved across the chromosomes. The use of genomic sequencing data is essential for the enhancement of cytogenetic analyses.     

Aluminum resistance mechanisms in oat (Avena sativa L.)

Background and aims

Enhanced aluminum (Al) resistance has been observed in dicots over-expressing enzymes involved in organic acid synthesis; however, this approach for improving Al resistance has not been investigated in monocots. Among the cereals, oat (Avena sativa L.) is considered to be Al resistant, but the basis of resistance is not known.

Methods

A hydroponic assay and hematoxylin staining for Al accumulation in roots were used to evaluate Al resistance in 15 oat cultivars. Malate and citrate release from roots was measured over a 24?h period. A malate dehydrogenase gene, neMDH, from alfalfa (Medicago sativa L.) was used to transform oat.

Results

Oat seedlings were highly resistant to Al, as a concentration of 325?μM AlK(SO₄)₂ was needed to cause a 50% decrease in root growth. Most oat cultivars tested are naturally resistant to high concentrations of Al and effectively excluded Al from roots. Al-dependent release of malate and Al-independent release of citrate was observed. Al resistance was enhanced in a transgenic oat line with the highest accumulation of neMDH protein. However, overall root growth of this line was reduced and expression of neMDH in transgenic oat did not enhance malate secretion.

Conclusions

Release of malate from oat roots was associated with Al resistance, which suggests that malate plays a role in Al resistance of oat. Over-expression of alfalfa neMDH enhanced Al resistance in some lines but was not effective alone for crop improvement.     

Phytotoxic substances with allelopathic activity may be central to the strong invasive potential of Brachiaria brizantha

The grass Brachiaria brizantha, native to eastern Africa, becomes naturalized and dominant quickly in the non-native areas. It was hypothesized that phytotoxic chemical interaction between this plant and native plants may play an important role in the invasion of B. brizantha. However, no potent phytotoxic substance has been reported in this species. Therefore, we investigated possible allelopathic activity and searched for phytotoxic substances with allelopathic activity in B. brizantha. An aqueous methanol extract of B. brizantha inhibited the growth of roots and shoots of garden cress (Lepidium sativum), lettuce (Lactuca sativa), timothy (Phleum pratense) and ryegrass (Lolium multiflorum) seedlings. The extract was purified by several chromatographic runs and three allelopathically active substances were isolated and identified by spectral analysis as (6R,9R)-3-oxo-α-ionol, (6R,9S)-3-oxo-α-ionol and 4-ketopinoresinol. (6R,9R)-3-Oxo-α-ionol and (6R,9S)-3-oxo-α-ionol inhibited root and shoot growth of garden cress at concentrations greater than 30 and 10 μM, respectively. The activity of (6R,9S)-3-oxo-α-ionol was 5.3- to 6.2-fold that of (6R,9R)-3-oxo-α-ionol. The stereochemistry of the hydroxyl group at position C-9 may be important for the inhibitory activities of those compounds. 4-Ketopinoresinol inhibited root and shoot growth of garden cress at concentrations greater than 30 μM. The growth inhibitory activity of (6R,9S)-3-oxo-α-ionol was the greatest and followed by 4-ketopinoresinol and (6R,9R)-3-oxo-α-ionol. These results suggest that those phytotoxic substances may contribute to the allelopathic effect caused by B. brizantha and may be involved in the invasion of B. brizantha.     

Population of Scaptocoris castanea Perty (Hemiptera: Cydnidae) in a Crop–Livestock Integration System

The population of the brown root stink bug, Scaptocoris castanea Perty, was studied in a crop?Clivestock integration system. This integrated system consisted of corn crop associated with Brachiaria decumbens pasture at the Unidade de Pesquisa e Desenvolvimento, in S?o José do Rio Preto (49°26???W; 20°49???S), state of S?o Paulo, Brazil. In each plot of the integrated crop?Clivestock system, which was consisted of six treatments and four replicates, four 0.25?m² and 0.30-m-deep samples of soil and roots were taken for analysis. These samples were stratified in 0.10-m layers, in which the number of nymphs and adults of S. castanea were assessed. These evaluations occurred monthly, from November 2008 to April 2009. The number of brown bug nymphs and adults was higher in areas where corn was cultivated for two consecutive years and in plots where pasture was renewed every 2?years. Lower insect population densities were observed in plots that remained as Brachiaria pasture. Therefore, S. castanea population is larger in integrated crop?Clivestock system (corn crop associated with B. decumbens), with no tillage.     

Isolation and characterization of microsatellite markers for Brachiaria brizantha (Hochst. ex A. Rich.) Stap

The first set of nuclear simple sequence repeat (SSR) loci for Brachiaria brizantha (Hochst. ex A. Rich.) Stap is described. A microsatellite-enriched library was constructed and 19 loci were characterized. About 13 SSR loci were found to be polymorphic and across-taxa amplification tests showed that six of them can be transferred to four other species of Brachiaria. This new SSR resource will be a powerful tool for population genetic studies of B. brizantha, for interspecific genetic studies within the genus Brachiaria, for mapping and for marker assisted selection in breeding.     

Belowground ABA boosts aboveground production of DIMBOA and primes induction of chlorogenic acid in maize

Plants are important mediators between above- and belowground herbivores. Consequently, interactions between root and shoot defenses can have far-reaching impacts on entire food webs. We recently reported that infestation of maize roots by larvae of the beetle Diabrotica virgifera virgifera induced shoot resistance against herbivores and pathogens. Root herbivory also enhanced aboveground DIMBOA and primed for enhanced induction of chlorogenic acid, two secondary metabolites that have been associated with plant stress resistance. Interestingly, the plant hormone abscisic acid (ABA) emerged as a putative long-distance signal in the regulation of these systemic defenses. In this addendum, we have investigated the role of root-derived ABA in aboveground regulation of DIMBOA and the phenolic compounds chlorogenic acid, caffeic and ferulic acid. Furthermore, we discuss the relevance of ABA in relation to defense against the leaf herbivore Spodoptera littoralis. Soil-drench treatment with ABA mimicked root herbivore-induced accumulation of DIMBOA in the leaves. Similarly, ABA mimicked aboveground priming of chlorogenic acid production, causing augmented induction of this compound after subsequent shoot attack by S. littoralis caterpillars. These findings confirm our notion that ABA acts as an important signal in the regulation of aboveground defenses during belowground herbivory. However, based on our previous finding that ABA alone is not sufficient to trigger aboveground resistance against S. littoralis caterpillars, our results also suggest that the ABA-inducible effects on DIMBOA and chlorogenic acid are not solely responsible for root herbivore-induced resistance against S. littoralis.Key words: induced resistance, Spodoptera littoralis, Zea mays, Diabrotica virgifera, DIMBOA, chlorogenic acid, absisic acid, priming     

Responses to drought of five Brachiaria species. II. Water relations and leaf gas exchange

Neotropical savannas are exposed to recurrent dry periods of varied duration, and forage grasses must be able to cope with such temporal stresses to maintain productive pastures. This study compared leaf water relations and net photosynthesis under drought of five perennial Brachiaria species: the tufted B. brizantha (CIAT 6780), the semi-stoloniferous B. decumbens (CIAT 606) and B. mutica, and the stoloniferous B. humidicola (CIAT 679) and B. dictyoneura (CIAT 6133). Plants of the five grasses were grown in large pots and subjected to drought by suspending watering until first wilting symptoms (14 days for B. brizantha, B. decumbens and B. mutica, and 29 days for B. humidicola and B. dictyoneura). Afterwards, they were re-watered and a second soil dry cycle was imposed. Time trends in leaf water potential (_l), relative water content (RWC), osmotic potential at full turgor (₀ ¹⁰⁰), stomatal conductance (Gs) and net photosynthesis (A) of stressed (DT) plants were compared to those of well-irrigated (CT) plants. Predawn _l in DT plants decreased to a minimum of –1.5 and –2.0 MPa in B. brizantha and B. mutica, compared to –2.5 to –3.0 MPa in B. decumbens, B. humidicola and B. dictyoneura. RWC decreased up to 50% in B. brizantha, compared to 75% in the other species. In B. humidicola, B. dictyoneura and in a lesser extent, B. decumbens, leaves of DT plants adjusted osmotically, by an apparent accumulation of nutrient solutes, at a rather constant ratio of turgid to dry weight of the tissue. Calculated osmotic adjustment ranged between 0.38 (B. decumbens) to 0.87 MPa (B. humidicola). This adjustment in ₀ ¹⁰⁰ was in some cases maintained 7 days after re-watering. In B. brizantha and B. mutica, Gs and A were significantly affected by drought, with maximum reduction percentages at the second drought period of 65 and 80%, respectively. The corresponding reduction in B. decumbens was 53 and 55%, respectively; whereas in B. humidicola and B. dictyoneura Gs and A were reduced less than 20%. In all species, re-watering allowed for the water relations (except ₀ ¹⁰⁰) and photosynthetic activity of leaves of DT plants to reach values comparable to those of CT plants. Results are discussed in term of root morphology and soil water extraction pattern, as well as leaf traits that may contribute to withstand drought under moderate soil water stress.