Grass species can be classified into different functional types based on their growth strategies, and contrasting persistence strategies are observed in different grass species. Excluding seedling recruitments, changes in populations of grasses are basically a trade-off between natality and mortality of tillers. We hypothesised that the persistence pathway of perennial grasses is linked to their growth strategy, regardless whether they are growing as monoculture or as a mixture. Species with contrasting growth strategies (Arrhenatherum elatius L., Dactylis glomerata L., and Festuca arundinacea Schreb.) were cultivated as monocultures and as a mixture and their tiller natality and mortality were evaluated for two years after swards establishment. All pastures maintained their population size during the experimental period, although decreases in tiller densities occurred during the warmer season. Arrhenatherum elatius had the highest tiller mortality and natality whereas the F. arundinacea had the lowest ones. Arrhenatherum elatius had many tillers appearing in all seasons but their tillers were short-lived. Conversely, F. arundinacea and D. glomerata developed numerous tillers during autumn and winter and their tillers survived, on average, almost six and three times longer than those of A. elatius, respectively. There were no differences in tillering dynamics among populations grown in monocultures or in the mixture. Regardless of whether they were cultivated in monocultures or as a mixture, the persistence pathway of perennial grasses is linked with their growth strategies with exploitative species presenting a high tiller turnover throughout the year whereas the persistence of more conservative species is based on a high tiller survival.
Oat and ryegrass intercropping in pastures is widely used in regions with subtropical climates. The aim of this study was to evaluate the tiller size/density compensation mechanisms in monoculture and intercropping swards of black oats (Avena strigosa Schreb cv. IAPAR 61) and annual ryegrass (Lolium multiflorum Lam. cv common) under intermittent grazing. Treatments (black oat, annual ryegrass and their mixture) were assigned according to a complete randomized block design with four replicates. Ryegrass, oat and intercropped pastures were grazed when the swards reached a height of 17, 25 and 23 cm, respectively, and with a level of defoliation of 40%. The aerial biomass was determined with a rising plate meter, and the tiller population density (TPD) was estimated by counting tillers in three 10 cm diameter PVC rings per paddock. The mass per tiller was estimated based on the aerial biomass and the TPD of each paddock. Total herbage production did not differ among treatments, with values around 7400 kg DM ha−1. TPD decreased and mass per tiller increased linearly in the monoculture treatments. Tiller size/density compensation was observed in the three plant communities (treatments) according to the self‐thinning rule. In addition, no relationships were found when each species was analysed individually in the intercrop treatment. The results suggest that species in grass mixed swards adjust their population to keep a relatively constant leaf area index (LAI) over the grazing seasons, and that would help pastures to stabilize herbage production.
Multi-species pastures have been viewed as a means to increase forage production relative to monocultures through enhanced three-dimensional occupation of the canopy, which will intercept and use the incident radiation more efficiently. For this to occur, increased production of photosynthetically active tissues such as leaves is required. We tested the hypothesis that intercropping of black oats (Avena strigosa Schum.) and annual ryegrass (Lolium multiflorum Lam.) can increase total and leaf forage production compared with their monocultures. Monocultures and mixed pastures of black oat and annual ryegrass were established, and their tiller morphogenetic and structural traits, as well as components of herbage production, were measured throughout the usage period of the pastures. There was no difference between treatments for total growth rate (77.1 kg dry matter (DM) ha–1 day–1), but annual ryegrass pastures presented the greatest stem growth rate (38.1 kg DM ha–1 day–1). Taking into account only the vegetative phase, the intercropped pasture produced 20% more leaf than the monocultures. There were no differences between species, either in monoculture or intercropped, for phyllochron, final leaf length, leaf elongation duration and leaf elongation rate. The most important modification in morphogenetic variables due to the inter-specific competition was an increase in senescence rate of annual ryegrass leaves. The average specific green leaf weight in the intercropping pastures corresponded to 84.6% and 137.5% of those values observed for black oat and annual ryegrass pastures, respectively. On the other hand, the intercropping pastures presented 43.5% more tillers than the black oat pastures and 17.8% fewer tillers than the annual ryegrass pastures. It is suggested that intercropping black oat and annual ryegrass does not change tiller ontogenetic processes and that the association of their different size and shape in intercropped pastures could increase pasture leaf production over their monocultures.
Annual temperate grasses go through abrupt changes in morphological composition during the growing season, which can affect ingestive behavior of grazing cattle. However, few studies have examined the effects of gradual structural changes in annual temperate pastures on the herbage intake rate during the growing season. We hypothesized that lenient grazing (removal of 40% of the initial pre‐grazing height) maintains short‐term intake rate (STIR) of grazing cattle over the entire growing season, even with decreased leaf/stem ratios. The studied pastures contained Avena strigosa, Lolium multiflorum and a mixture of the two species. Short‐term intake rate was measured using the double weighing technique, evaluating both bite rate and bite mass. The percentage of stem + pseudostem and leaf lamina lengths in the herbage mass were measured. Relationships between sward structure and animal ingestive behavior were also examined. As the pasture growing season progressed, the STIR of heifers grazing annual temperate grass swards was affected more by the time required to take a bite than by bite mass. Moreover, the sward structure also changed after each grazing cycle, reducing the pre‐grazing leaf/stem ratio. However, reductions in the pre‐grazing leaf lamina mass (up to 37% of the sward herbage mass) and/or in the leaf lamina length (up to 52% of the extended tiller height) did not affect the STIR of grazing heifers, since the animals maintained their bite mass and time per bite. In summary, with a grazing down target of 40% of the pre‐grazing height, reductions in STIR of heifers grazing annual temperate grasses occurred when the pre‐grazing leaf lamina percentage was <37% of the sward herbage mass and/or <52% of the extended tiller height. These results suggested that to avoid STIR reductions during the last grazing cycles, managements to stimulate tillering and delay stem + pseudostem elongation are necessary and should be investigated.
SUMMARYMixing species with different tillering peaks may enhance pasture stability, but intercropping may also alter the plants’ tillering patterns. This study aimed to evaluate tillering dynamics in black oat (Avena strigosa Schreb.) and annual ryegrass (Lolium multiflorum Lam.) pastures grown as monocultures or intercropped. Three following treatments are established: black oat and annual ryegrass pastures grown as monocultures and an intercrop composed of these two species. Tillering dynamics were measured for black oat and annual ryegrass populations separately. When intercropped, tiller birth rates of black oat decreased (20.0 vs. 28.9 tillers 100 tillers−1) and those of annual ryegrass increased (30.5 vs. 14.3 tillers 100 tillers−1), compared to their monocultures. Tiller death rates for annual ryegrass did not differ between monoculture and intercropping (23.9 tillers 100 tillers−1), but black oat presented higher mortality in monocultures (48.8 vs. 36.4 tillers 100 tillers−1). The black oat monoculture had the lowest population stability index (0.80), whereas annual ryegrass in monoculture and intercropped pastures exhibited greater values (on average, 0.92). Our results indicated that black oat and annual ryegrass present distinct tillering dynamics whether grown as monoculture or intercropped, and suggest that intercropping species with elevated death rates (black oat) with later species (annual ryegrass) could be an important tool for maintaining pasture stability throughout the growing season.
Core Ideas Mixing grass forage species can increase forage yield and stability. Overyielding was more influenced by increasing leaf biomass rather than stem biomass. Dominant grasses with contrasting growth strategies can compose productive and stable pastures. Mixtures composed of species with different growth strategies (e.g., exploitative and conservative), which have different abilities for resource capture and use, are thought to improve grassland ecosystem services. However, plant species that compete better for limiting resources can become dominant. This study examined whether potential dominant perennial grasses with different growth strategies can compose productive and stable forage mixtures under low competition for resources (nutrients and light). Arrhenatherum elatius L. (exploitative), Festuca arundinacea Schreb. (conservative), and Dactylis glomerata L. (moderately exploitative) were sowed as monocultures and as a three‐species mixture. Pastures were fertilized to maintain high soil fertility levels and mowed by half when their canopies reached 20 cm in height (95% of light interception in the vegetative stage) allowing for over 2 yr of data collection. The proportion of each species in the mixture remained unaltered throughout the experimental period. Transgressive overyielding was not observed; however, the mixture presented a similar yield to the most productive monocultures (10,200 kg DM ha−1 yr−1), although D. glomerata, which presented the lowest yield in monoculture (6400 kg DM ha−1 yr−1), comprised 68.1% of post‐cutting biomass in the mixture. Arrhenatherum elatius and F. arundinacea populations presented less yield variance in monoculture, but higher yield stability was observed in the mixture. Overyielding in the three studied species was strongly affected by leaf production. Mixtures composed of perennial grasses with different growth strategies when grown in nutrient‐rich soil and under conditions of low light competition can compose productive and stable swards, while maximizing leaf production.
RESUMOPastos consorciados de aveia e azevém anual têm sido recomendados para uso no inverno/primavera em ambientes subtropicais. No entanto, o impacto do pastejo leniente sobre parâmetros qualitativos e produtivos destes, comparativamente aos seus monocultivos, é pouco conhecido. Objetivou-se avaliar possíveis variações nas composições morfológica e bromatológica, além da produção de biomassa em pastos de aveia-preta (Avena strigosa cv. IAPAR 61) e azevém anual (Lolium multiflorum Lam. cv. comum) quando cultivados puros ou em consórcio ao longo de toda estação de crescimento e submetidos a pastejo leniente. Os três tratamentos (aveia pura, azevém puro e aveia + azevém em consórcio) foram distribuídos em um delineamento em blocos completos ao acaso, com quatro repetições. Novilhas da raça Holandesa foram utilizadas como agentes de desfolhação, e os pastos foram rebaixados em 40% da altura inicial quando as alturas atingiam 17, 20 e 23cm para o azevém, o consórcio e a aveia, respectivamente. A relação lâmina:colmo diminuiu com o avanço da estação de crescimento em todos os tratamentos, mas a proporção de colmos no estrato pastejável se manteve abaixo de 20% da matéria seca até o início do estádio de desenvolvimento reprodutivo, independentemente do tipo de pasto. Os teores de proteína bruta e fibra em detergente neutro no estrato pastejável foram, respectivamente, superiores a 16% e inferiores a 50% nos quatro primeiros ciclos de pastejo. O consórcio e o azevém puro apresentaram as maiores taxas de acúmulo médio de forragem (32,6kg MS ha -1 dia -1 ), porém o maior período de utilização do consórcio resultou em maior produção total de matéria seca (4809kg MS ha -1 ). O pastejo leniente não prejudica a qualidade da forragem no estrato pastejável durante todo o estádio de desenvolvimento vegetativo de pastos anuais de inverno. A associação entre aveia-preta e azevém anual aumenta a produção de matéria seca total de forragem em comparação aos mesmos pastos cultivados puros.
Background and aims: The concepts of critical nitrogen (N) concentration and nitrogen nutrition index (NNI) provide effective tools for assessing the N nutritional status of pastoral systems. However, no conclusive data exist concerning the effects of stubble height at harvest on the N dilution curve [N concentration (N%) = a(biomass)−b] and mixed species cultivation on N nutritional status of forage plants with contrasting growth strategies. To evaluate these effects, three cool‐season perennial grasses (exploitative, Arrhenatherum elatius L.; moderately exploitative, Dactylis glomerata L.; conservative, Festuca arundinacea Scherb.) were grown under different cultivation methods either as pure stands or as a mixture and conducted two experiments. Methods: In experiment I, canopies of different grass species pure stands and mixtures were subjected to defoliation managements aimed to not exceed the critical LAI at harvest. Forage samples were collected from October 2015 to August 2016 to determine the N nutritional status throughout the seasons. In experiment II, half of each pure stand was mowed to 10 cm in height, and the other half to ground level. Thereafter, herbage samples were taken during regrowth to determine N dilution curves in both scenarios (from ground level and from stubble height of 10 cm). Results: We verified an NNI above 0.8 during autumn and winter for A. elatius, D. glomerata, and the mixture but only in winter for F. arundinacea. Dactylis glomerata grown in the mixture had a lower N% than its pure stand during autumn and winter, whereas A. elatius and F. arundinacea showed no differences irrespective of cultivation method throughout the year. The three forage species presented classical dilution curves when regrowth was initiated from ground level, but quadratic trends were observed when regrowth was initiated from stubble heights of 10 cm. Conclusions: Overall the N nutritional status of cool‐season perennial grasses was not affected by cultivation method when pastures were managed in conditions of non‐limiting N supply and with pre‐cut/grazing height targets derived from critical LAI measured at the full vegetative growth stage. Moreover, the N nutrition status in pastures managed under moderate/lenient defoliation should not be evaluated during periods of less intense canopy renewal and/or right after the cut/grazing events.
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