Phylogenomic analysis reveals five independently evolved African forage grass clades in the genus Urochloa.

BACKGROUND AND AIMS: The grass genus Urochloa (Brachiaria) sensu lato includes forage crops that are important for beef and dairy industries in tropical and sub-tropical Africa, South America and Oceania/Australia. Economically important species include U. brizantha, U. decumbens, U. humidicola, U. mutica, U. arrecta, U. trichopus, U. mosambicensis and Megathyrsus maximus, all native to the African continent. Perennial growth habits, large, fast growing palatable leaves, intra- and interspecific morphological variability, apomictic reproductive systems and frequent polyploidy are widely shared within the genus. The combination of these traits probably favoured the selection for forage domestication and weediness, but trait emergence across Urochloa cannot be modelled, as a robust phylogenetic assessment of the genus has not been conducted. We aim to produce a phylogeny for Urochloa that includes all important forage species, and identify their closest wild relatives (crop wild relatives). Finally, we will use our phylogeny and available trait data to infer the ancestral states of important forage traits across Urochloa s.l. and model the evolution of forage syndromes across the genus. METHODS: Using a target enrichment sequencing approach (Angiosperm 353), we inferred a species-level phylogeny for Urochloa s.l., encompassing 54 species (~40 % of the genus) and outgroups. Phylogenies were inferred using a multispecies coalescent model and maximum likelihood method. We determined the phylogenetic placement of agriculturally important species and identified their closest wild relatives, or crop wild relatives, based on well-supported monophyly. Further, we mapped key traits associated with Urochloa forage crops to the species tree and estimated ancestral states for forage traits along branch lengths for continuous traits and at ancestral nodes in discrete traits. KEY RESULTS: Agricultural species belong to five independent clades, including U. brizantha and U. decumbens lying in a previously defined species complex. Crop wild relatives were identified for these clades supporting previous sub-generic groupings in Urochloa based on morphology. Using ancestral trait estimation models, we find that five morphological traits that correlate with forage potential (perennial growth habits, culm height, leaf size, a winged rachis and large seeds) independently evolved in forage clades. CONCLUSIONS: Urochloa s.l. is a highly diverse genus that contains numerous species with agricultural potential, including crop wild relatives that are currently underexploited. All forage species and their crop wild relatives naturally occur on the African continent and their conservation across their native distributions is essential. Genomic and phenotypic diversity in forage clade species and their wild relatives need to be better assessed both to develop conservation strategies and to exploit the diversity in the genus for improved sustainability in Urochloa cultivar production.

The use of light-transparent acetate sheet to measure canopy height of marandu palisadegrass: the influence of development stage and sward height

The use of light-transparent acetate sheet (LAS) may generate variations in the height measured in the sward. However, we compared the use or non-use of LAS to measure the average sward height of marandu palisadegrass (Urochloa brizantha syn. Brachiaria brizantha cv. Marandu) pastures. In the first experiment, the methods were applied in pastures with different average heights (15, 25, 35, and 45 cm), managed under continuous stocking with sheep, and with predominance of vegetative tillers. The shorter pastures (15 and 25 cm) had similar heights with or without the use of the LAS. However, in the taller pastures (35 and 45 cm), the use of LAS resulted in lower height values, compared with its absence. In the second experiment, we compared both methods for measuring the height of deferred marandu palisadegrass and fertilized with two nitrogen levels (50 and 200 kg ha-1) and with high presence of reproductive tillers. A completely randomized design with four replicates was adopted. With 200 kg ha-1 of N, the canopy height was greater than when it was fertilized with 50 kg ha-1 of N. With the use of LAS, the height of deferred and reproductive canopy was greater than without it. In a taller marandu palisadegrass pasture with predominance of vegetative tillers, the use of LAS results in smaller height, contrary to what occurs in deferred swards with reproductive tillers.

The sward height at the early stockpiling period modifies the structural characteristics of marandu palisadegrass under continuous stocking

The control of sward height at the early stockpiling period influences the structural characteristics of stockpiled sward in winter. So, we conducted this work on the Cerrado region of Brazil based on the hypothesis that the reduction of pasture height at the beginning of stockpiling period results in: (i) lower stock of forage mass; and (ii) higher live leaf mass, but lower stem mass in stockpiled pasture during the winter. The treatments were four sward heights (15, 25, 35, and 45 cm) at early stockpiling period of marandu palisadegrass (Brachiaria brizantha syn. Urochloa brizantha cv. Marandu). The experiment was conducted in a completely randomized design, with three replications (paddocks with 800 m²). The stockpiled swards were evaluated during the grazing period in winter on days 7 (beginning), 45 (middle) and 90 (end). The vegetative tiller number (VEGT), live leaf mass (LLM) and leaf area index (LAI) were greater in the 15 cm stockpiled pasture than the other treatments. Greater heights (35 and 45 cm) at stockpiling resulted in greater forage mass than lower heights (15 and 25 cm). The VEGT, LLM, number of reproductive tillers, live stem mass, forage mass and LAI values were higher at the beginning than at the end of the grazing period. Our hypothesis has been proven, so that the reduction of sward height of the marandu palisadegrass at the beginning of the stockpiling period results in lesser forage mass, but with better structural characteristics.

How does the initial sward height and the grazing period influence the spatial variability of vegetation in deferred signal grass pastures?

The objective of this study was to evaluate the spatial variability of structural characteristics of deferred pastures of Brachiaria decumbens cv. Basilisk (signal grass) subjected to associations of sward heights (10, 20, 30, and 40 cm) at the beginning of deferment and grazing periods (1, 28, 56, 85, and 113 days). The experiment was arranged in a split-plot and completely randomized-block design with two replicates. Pastures remained deferred from March to June 2010. From June to the beginning of October 2010, deferred pastures were utilized by steers under continuous grazing and at a fixed initial stocking rate of 3.5 AU ha-1. At the beginning of the deferment, the coefficient of variation (CV) for pasture height was reduced linearly with the pasture height. During the grazing period, in the winter, the CV for pasture height was not influenced by initial pasture height; however, it responded quadratically to the grazing period, with a maximum value of 36.4% at 71 days of the period of utilization. The CV for tiller height was reduced linearly with pasture height at the beginning of deferment but increased linearly along with the grazing period. The initial pasture height did not change the CV of the falling index. However, the latter was quadratically influenced by the grazing period, with a maximum value of 59.5% at 67 days of the utilization period. Signal grass pastures deferred at a lower height have a higher spatial variability of the vegetation. In the grazing period, there are changes in the spatial variability of the vegetation of the deferred signal grass.

Does amount of cut vegetal residue on the canopy influence growth and the structural characteristics of marandu palisadegrass?

The pasture mowing in late winter removes the old forage, improving the pasture structure in spring and summer. However, the residue after mowing can affect tillering, thus limiting forage production and the structural characteristics of pasture. We hypothesized that the high amount of cut vegetal residues on the plants in late winter causes a decrease in forage production and modifies the structural characteristics of the forage canopy during the spring and summer. The treatments were four cut vegetal residues (0; 2,000; 4,000 and 8,000 t ha-1 of natural material) deposited on Urochloa brizantha cv. Marandu canopy in late winter. After that, the forage production and structure characteristics were evaluated during the spring and summer for two years. The complete randomized block design, with four replications, was used. The defoliation management was characterized by the adoption of pre- and post-cut heights of 25 and 15 cm, respectively. For both years, there was no effect of cut vegetal residue on all the characteristics evaluated. No variable was affected by the interaction cut vegetal residue amount × year of evaluation. The dead leaf blade percentage was greater in year 2 (6.9%) than in year 1 (3.5%). The average values of variables were live leaf blade percentage = 84.2%; live stem percentage = 8.7%; dead stem percentage = 1.9%; number of vegetative tiller = 653 tiller/m2; number of reproductive tiller = 5 tiller m-2. The forage production rate presented an average value of 46 kg/ha/day of dry matter. The cut vegetal residue amount of up to 8,000 t/ha of natural material on the plants in late winter does not affect the forage production or modifies the structural characteristics of marandu palisadegrass during the subsequent spring and summer.

Morphogenetic, structural and production traits of marandu grass under nitrogen rates in Neo soil / [Características morfogênicas, estruturais e produtivas do capim-marandu sob doses de nitrogênio em Neossolo]

Most sandy soils have low natural fertility and low levels of organic matter, making nitrogen (N) fertilization essential. Thus, five doses of N were applied (0, 75, 125, 175 and 225mg dm-³) in a randomized block design to evaluate the effects of nitrogen fertilization on the morphogenetic, structural and production characteristics of Brachiaria brizantha cv. Marandu in a Quartzarenic neosoil. The doses of N did not affect the height of the canopy. The leaf elongation rate, final leaf length and number of live leaves increased linearly at the doses of N. Leaf appearance rate, stem elongation rate, leaf lifespan, phyllochron, leaf senescence rate and tiller density showed a quadratic response to the rates There was also an effect of N rates in herbage mass, leaf mass, stem mass, which increased linearly. Brachiaria brizantha cv. Marandu cultivated in Quartzarenic neosoil requires higher doses of N, 175 and 225mg dm-³. Under these conditions, increases in its morphogenetic, structural and productive characteristics are observed. These findings may not be repeated in the most fertile soils with the greatest capacity to supply N.(AU)

A maioria dos solos arenosos tem baixa fertilidade natural e baixos teores de matéria orgânica, tornando a adubação com nitrogênio (N) essencial. Assim, foram aplicadas cinco doses de N (0, 75, 125, 175 e 225mg dm-³) em delineamento de blocos ao acaso, para se avaliarem os efeitos da adubação com nitrogênio nas características morfogênicas, estruturais e produtivas da Brachiaria brizantha cv. Marandu, em um Neossolo Quartzarênico. As doses de N não afetaram a altura do dossel. A taxa de alongamento foliar, o comprimento final da folha e o número de folhas vivas aumentaram linearmente em função das doses de N. A taxa de aparecimento de folhas, a taxa de alongamento do caule, o tempo de vida da folha, o filocrono, a taxa de senescência foliar e a densidade de perfilhos apresentaram resposta quadrática às doses de N. A massa da forragem, a massa foliar e a massa do caule aumentaram linearmente. Brachiaria brizantha cv. Marandu cultivada em Neossolo Quartzarênico requer maiores doses de N, 175 e 225mg dm-³. Nessas condições, são observados aumentos em suas características morfogenéticas, estruturais e produtivas. Esses achados podem não se repetir nos solos mais férteis e com maior capacidade de suprir N.(AU)

Características estruturais e produção de forragem do capim-piatã submetido a regimes de desfolhação intermitente / [Structural characteristics and forage production of piata grass submitted to intermittent defoliation management]

Objetivou-se avaliar as características estruturais e a produção de forragem do capim-piatã submetido à combinação de duas frequências de corte (95% e máxima interceptação de luz pelo dossel forrageiro, ILmax) e duas alturas pós-corte (15 e 20cm), no período de janeiro de 2012 a março de 2013. O delineamento utilizado foi em blocos ao acaso, com três repetições, em esquema fatorial 2x2, totalizando 12 unidades experimentais de 14m2 cada. Foram avaliadas as seguintes variáveis-respostas: altura do dossel no pré-corte, intervalo de corte, índice de área foliar pré e pós-corte, taxa de produção de forragem e as percentagens de colmo, folha e forragem morta na forragem produzida. A altura pré-corte decresceu durante o experimento (de 65,81 para 34,03cm), sendo a menor altura observada no dossel sob alta frequência de desfolhação (95% de IL). A taxa de produção de forragem foi maior nas épocas favoráveis ao crescimento da gramínea forrageira. As percentagens de folha, colmo e forragem morta foram semelhantes entre as frequências de corte. Assim, o capim-piatã sob manejo de desfolhação intermitente pode ser colhido com qualquer combinação de frequências de corte de 95% e ou máxima interceptação de luz, e alturas pós-corte de 15 ou 20cm.(AU)

The objective of this study was to evaluate the structural characteristics and forage production of the piatã grass submitted to the combination of two cutting frequencies (95% and maximum interception of light by the forage canopy, ILmax) and two post cutting heights (15 and 20cm) in the period from January 2012 to March 2013. The design was a randomized complete block with three replications, in a 2x2 factorial scheme, totaling 12 experimental units of 14m2 each. The following variables were evaluated: pre-cut canopy height, cut interval, pre and post-cut leaf area index, forage yield rate, and percentage of stem, leaf and dead forage in the forage produced. The pre-cut height decreased during the experiment (from 65.81 to 34.03cm), with the lowest height observed in the canopy under high frequency of defoliation (95% IL). The rate of forage production was higher in the seasons favorable to forage grass growth. The percentages of leaf, stem and dead fodder were similar between cut-off frequencies. Thus, piatã grass under intermittent defoliation management can be harvested with any combination of 95% and maximum light interception cut frequencies, and post cutting heights of 15 or 20cm.(AU)

Regrowth age modifies the leaf anatomy of Brachiaria genotypes / A idade de regeneração modifica a anatomia foliar dos genótipos de Brachiaria

Changes in leaf anatomy were evaluated in genotypes of Brachiaria brizantha, Brachiaria decumbens, Brachiaria ruziziensis and three Brachiaria ruziziensis clones through tissue proportion in internerval and midrib regions at three regrowth ages. Plants were grown and cutting was performed after 60 days. Further, leaves were sampled at 8, 15 and 29 regrowth days and processed with usual plant microtechnique. The internerval region showed higher parenchyma percentage at 15 days for Clone 95 and similar values at 15 and 29 days for Clone 1. The proportion of vascular bundles was lower after 15 days in Clones 1 and 95 and 29 days in B. brizantha. In the midrib, the parenchyma proportion was higher at 29 days in B. brizantha and lower at 15 days in B. ruziziensis. The proportion of vascular bundles was higher at 8 days in B. decumbens, B. brizantha and Clone 1, and lower at 29 days for Clones 97 and 95 and at 8 days in B. ruziziensis. Therefore, the regrowth age modifies the percentage of leaf tissues in Brachiaria genotypes, in which the fibers and vascular bundles increase at 29 days and 8-day-old leaves are not fully developed.

A anatomia foliar foi avaliada em genótipos de Brachiaria brizantha, Brachiaria decumbens, Brachiaria ruziziensis e três clones de Brachiaria ruziziensis; quanto à proporção de tecidos foliares, em três idades de rebrota. As plantas foram cultivadas e cortadas após 60 dias. As folhas foram amostradas aos 8, 15, 29 dias de rebrota e analisadas pela metodologia de microtécnica vegetal. Na região internervural, a proporção de parênquima foi maior aos 15 dias para o Clone 95, e aos 15 e 29 dias para o Clone 1. Proporção de feixes vasculares foi menor aos 15 dias para os Clones 1 e 95 e aos 29 dias para genótipo B. brizantha. Na nervura central, a proporção de parênquima foi maior aos 29 dias para B. brizantha e menor, aos 15 dias, para B. ruziziensis. Proporção dos feixes vasculares foi maior aos 8 dias para genótipo B. decumbens, B. brizantha e para o Clone 1, e menor aos 29 dias para os Clones 97 e 95; aos 8 dias, para genótipo B. ruziziensis. Portanto, a idade de rebrota modifica a porcentagem de tecidos foliares em genótipos de Brachiaria; contudo, fibras e feixes vasculares aumentam aos 29 dias; aos 8 dias, as folhas não estão totalmente desenvolvidas.

Genetic diversity and population structure analysis of the tropical pasture grass Brachiaria humidicola based on microsatellites, cytogenetics, morphological traits, and geographical origin.

Brachiaria humidicola (Rendle) Schweick. is a warm-season grass commonly used as forage in the tropics. Accessions of this species were collected in eastern Africa and massively introduced into South America in the 1980s. Several of these accessions form a germplasm collection at the Brazilian Agricultural Research Corporation. However, apomixis, ploidy, and limited knowledge of the genetic basis of this germplasm collection have constrained breeding activities. The objectives of this work were to identify genetic variability in the Brazilian B. humidicola germplasm collection using microsatellite markers and to compare the results with information on the following: (1) collection sites of the accessions; (2) reproductive mode and ploidy levels; and (3) genetic diversity revealed by morphological traits. The evaluated germplasm population is highly structured into four major groups. The sole sexual accession did not group with any of the clusters. Genetic dissimilarities did not correlate with either geographic distances or genetic distances inferred from morphological descriptors. Additionally, the genetic structure identified in this collection did not correspond to differences in ploidy level. Alleles exclusive to either sexual or apomictic accessions were identified, suggesting that further evaluation of the association of these loci with apospory should be carried out.

Phytoremediation in the tropics--influence of heavy crude oil on root morphological characteristics of graminoids.

When studying species for phytoremediation of petroleum-contaminated soils, one of the main traits is the root zone where enhanced petroleum degradation takes place. Root morphological characteristics of three tropical graminoids were studied. Specific root length (SRL), surface area, volume and average root diameter (ARD) of plants grown in crude oil-contaminated and uncontaminated soil were compared. Brachiaria brizantha and Cyperus aggregatus showed coarser roots in polluted soil compared to the control as expressed in an increased ARD. B. brizantha had a significantly larger specific root surface area in contaminated soil. Additionally, a shift of SRL and surface area per diameter class towards higher diameters was found. Oil contamination also caused a significantly smaller SRL and surface area in the finest diameter class of C. aggregatus. The root structure of Eleusine indica was not significantly affected by crude oil. Higher specific root surface area was related to higher degradation of petroleum hydrocarbons found in previous studies.