Dendrometric analysis
The absence of effect regarding the cultivation system on plant height can be explained by the greater intraspecific competition for light between plants that occurred in the monocrop system (5,000 plants ha− 1), since after 38 months of age, the plants had already reached their full development (4.5 m). Furthermore, M. caesalpiniifolia has rapid initial growth, which is one of the main advantages of this species in pasture areas, when compared to other arboreal and exotic legumes (Silva et al. 2014; Herrera et al. 2021). Some morphological responses under competition may occur in the crown area, branches, and stem shape, affecting wood production (Yang et al. 2019). For example, under competition for light, plants can promote the growth of supporting structures such as stems and branches, aiming to allow the leaves to reach the light (Cardwell, 1987). Furthermore, to intercept more light, plants change the balance between vertical and lateral growth, which eventually triggers architectural transformation (Clark, 2010).
This may explain the results of other studies performed in the same experimental area in the initial growth phase of M. caesalpiniifolia, for example, Carvalho et al. (2022) evaluated plant height in initial growth (29 months) and observed a significant difference (P < 0.05) between cultivation systems, with average plant height values of 2.4 m under monocrop, and 2.8 m in the silvopasture. Oliveira Neto (2022) evaluated M. caesalpiniifolia until 38 months of age and also recorded a significant effect, with higher plants (3.6 m) in the monocrop systems, compared to the silvopasture (3.4 m). Thus, the cultivation system may influence M. caesalpiniifolia height in its initial growth (38 months). When the plants reached their full development (38 to 60 months), in the case of the present study, no difference in plant height was observed between cultivation systems. According to Lorenzi (2002), M. caesalpiniifolia can vary in height between 5 and 8 m.
Herrera et al. (2021) evaluated the growth dynamics and nutritional value of different legume species gliricidia (Gliricidia sepium) and M. caesalpiniifolia in silvopasture system with signal grass (Urochloa decumbens) and observed a higher plants of M. caesalpiniifolia (6.1 m) compared to gliricidia (4.9 m), at seven years of age. The rapid growth of M. caesalpiniifolia in silvopasture when compared to gliricidia can be justified by its reduced consumption by grazing animals due to the presence of prickles and condensed tannins, which limit the animals' intake of this plant (Carvalho et al. 2022; Oliveira Neto, 2022). The opposite occurs with gliricidia due to its high acceptability, animals consume more of this legume ending up reducing their growth and development under grazed areas such as silvopasture (Herrera et al. 2021).
Concerning the forage mass, the absence of effect of the cultivation system (p > 0.05) can be explained due to the greater density in monoculture (5,000 plants ha− 1) compared to SSP (600 plants ha− 1). In denser cultivations, greater biomass production is expected, as well as greater production of wood per area. However, this condition in the monocrop system resulted in greater shading (trees with an average of 4.9 m) in the lower stratum (1.5 m) due to the elongation of the tree stem in this condition (Paciullo et al. 2011; Taiz et al. 2017). Forage production per plant (7 g DM plant− 1) and the forage mass were reduced due to the low incidence of light in the lower stratum of the M. caesalpiniifolia under monocrop. Wang et al. (2012) indicated that strong competition reduces available resources and decreases tree growth and biomass accumulation.
On the other hand, in the silvopasture system, the lower density resulted in less competition between plants contributing to higher production per plant (36 g DM plant− 1) (Table 1) when compared to the monocrop system of M. caesalpiniifolia. The reduced production per plant contributed to the reduction of forage mass (35 kg DM ha− 1) in the monocrop system, this may have contributed to minimizing the significant effect of the cultivation systems on the forage mass. Yang et al. (2019) reported that trees compete for both light and growing space; under strong competition, oak trees (Quercus liaotungensis) reduced the biomass of their branches to allow the biomass of the underground parts to develop, while at the same time prioritizing stem growth. Oliveira Neto (2022), evaluating forage mass up to 1.5m in height in the same experimental area between 2019–2020 and 2020–2021 in the dry period, reported forage mass values between 24 and 332 kg DM ha− 1, respectively.
The forage mass obtained in the present study for the monocrop and silvopasture are higher than those found by Silva et al. (2014) in areas with a density of 600 plants ha− 1, when harvesting branches and leaves at 54 months of age, they reported an average value of 6.49 kg DM ha− 1. On the other hand, Costa et al. (2016) and Oliveira et al. (2018) reported average values ranging from 94 to 624 kg DM ha− 1 in areas intercropped with M. caesalpiniifolia and signal grass during the period from September 2012 to August 2013, with a density of 2,500 plants ha− 1, and harvesting at a height up to 1.5 m. Forage mass reductions in legume trees were also reported by Herrera et al. (2021) in silvopasture with grass, these authors reported that water availability at the time of harvesting was a factor of influence.
This was probably due to the greater number of stems in monocrop trees compared to silvopasture. Apolinário et al. (2015) reported that the development of the diameter at breast height of M. caesalpiniifolia was inversely related to the increase in the number of stems. This plant has the characteristic of dichotomous branches that are mobilized to stimulate stem elongation and growth in monoculture, however, this is a characteristic of the plants that etiolate, concentrating their reserves to lengthen the stems, consequently, they have a smaller basal diameter and diameter at breast height (Herrera et al. 2021).
M. caesalpiniifolia is a species that has dense and compact wood, with low moisture content. The commercial value of M. caesalpiniifolia wood, with a diameter between 12.6–21.9 cm in circumference, is estimated approximately between US$ 0.9 to 1.6 per stem-stake; and with a diameter of 25.5 cm between US$ $1.9 to 2.4 stem-stake, both prices estimated for 2.2 m long stems. According to Herrera et al. (2021), the stems with these diameters correspond to intermediate and thick branches and have the potential to contribute to increasing the producer's income, especially in silvopasture system with M. caesalpiniifolia. The trees produced in the present study had their diameter at breast height within the expected commercial size and could contribute to improving produce incomes, especially with the plants with the largest diameter at breast height as observed for the M. caesalpiniifolia growing in the silvopasture. However, it is important to consider that since implementation, the trees have not undergone any thinning management actions. According to Murta Junior et al. (2020), thinning can stimulate the production of high-quality wood by obtaining large trees in a shorter time, altering the dendrometric characteristics of the remaining trees.
The basal diameter was larger (> 19%) in the silvopasture compared to the observed in the monocrop system, probably due to less intraspecific competition for light, water, and nutrients (Silva et al. 2021a). Furthermore, this variable increased significantly throughout the evaluation period, with an average basal diameter of 18.7 cm during the experimental period. Herrera et al. (2021) and Carvalho et al. (2022) reported a basal diameter of 7.6 cm and 13.2 cm, at 84 and 29 months, respectively. In the present study, after 60 months of establishment, the basal diameter was 24.0 cm, representing 3.16 and 1.8 times greater than that reported by these authors, respectively. This probably occurred because this plant has an extensive root system that can access nutrients and water in deeper layers of the soil (Lira Jr. et al. 2020; Silva et al. 2021) and thus continued its growth. Furthermore, it is important to monitor the height of trees to improve the management of the forestry component, aiming to understand its potential in the environment in which it is located (Rajab-Pourrahmati et al. 2018).
The lower forage mass during the dry period can be justified because M. caesalpiniifolia is a deciduous species, with senescence of the leaves and detachment of branches during the dry period (Castro Filho et al. 2016). Although there was an increase in forage mass of 37% in the rainy season (Fig. 4B), the result obtained in the present study was lower than that reported by Oliveira Neto (2022) (661 kg DM ha− 1). The forage mass produced by M. caesalpiniifolia is not readily accessible for animal consumption due to its height. Senescing leaves may contribute to nutrient cycling via litter deposition and N cycling to the system, indirectly contributing to animal production (Apolinário et al. 2015; Figueiredo et al. 2023).
In the first year of evaluation in the monocrop system there was a higher forage mass (69 kg DM ha− 1) compared to the silvopasture systems in both years (22 kg DM ha− 1). This variation can be explained due to the greater number of plants in monoculture (5,000 plants ha− 1) compared to silvopasture (600 plants ha− 1). Furthermore, in the second year the reduction in forage mass occurred due to the shading caused by the growth of the plant stems in search of light, thus, the forage mass was limited in the lower stratum of the trees. Legume species produce less forage when compared to grasses due to their C3 metabolism, and the presence of photorespiration in high-temperature conditions (Taiz et al. 2017). It is worth mentioning that the production of M. caesalpiniifolia considered only thin branches up to 1.5 m in height.
Nutritional value
Greater dry matter levels during the dry period are normally observed, since the moisture levels in the soil and air are reduced, due to the absence of precipitation (Pereira et al. 2022). During this period, there is a restriction in plant growth (Mukarram et al. 2021), resulting in higher concentrations of DM in plant structures. The reduction in crude protein content during the dry period occurs due to the loss of quality, induced by the maturity of the leaves (Fig. 6A). Carvalho et al. (2022) observed a greater crude protein content in M. caesalpiniifolia leaves during the rainy season, with the content being reduced in the dry season with the advancement of leaf maturity and also due to deciduousness. Other studies (Bhardwaj et al. 2021; Navale et al. 2022; Sharma et al. 2023) reported variations in the crude protein content in the foliage of other tree species during different times of the year. Seasonal variations in temperature, precipitation, and other climatic conditions affect the photosynthetic process, which alters forage production and nutritional value (Ravhuhali et al. 2022). The crude protein content in the leaves is considered one of the most important nutritional parameters for animal health and productivity (Sharma et al. 2023), as ruminants need at least 70 g kg− 1 of crude protein to maintain the microbial metabolic processes in the rumen. Based on this, in the present study, the crude protein content in the rainy and dry seasons can meet the nutritional requirements for ruminants.
Plants generally have higher growth rates during the rainy period in the tropics, due to the greater availability of water and nutrients in the soil (Piao et al. 2019), resulting in greater biomass production and, consequently, greater accumulation of fibrous and support tissues, via increased thickness of cell walls (Ferreira et al. 2023). Several abiotic factors (e.g., precipitation, temperature, and soil fertility) also have a direct influence on the nutritional value of forage (Silva et al. 2021). Furthermore, the branches ≤ 5.0 mm sampled with the leaves may have contributed to increasing the ADF content in both periods of the year, a fact that also explains the low IVDMD coefficient observed. Herrera et al. (2021) evaluated the nutritional value of M. caesalpiniifolia in a silvopastoral system and reported an ADF content of 392 g kg− 1 DM in the leaves. Concerning the IVDMD, as leaf maturity advances, thickening and increased lignification of the cell wall occur, resulting in a decline in digestibility (Lima et al. 2019), a fact that probably occurred in the dry period, causing low IVDMD (248.98 g kg− 1 DM). Low digestibility rates can reduce forage consumption, due to the decrease in passage rate (Mertens, 2010), thus compromising animal performance over time. The variation in NDF was associated with the proportions of thin branches and leaves harvested in each sampling. According to Van Soest (1994), when NDF values exceed the range of 550–600 g kg− 1 DM, there is a negative effect on forage consumption and digestibility, consequently harming animal performance.
The CP levels obtained in the present study confirm the importance of using legumes in grazing production systems, as content above the 7% required for the maintenance of the ruminal microbiota was observed. In addition to increasing CP content in the diet of grazing animals, N-rich litter is capable of providing an additional source of this nutrient for the pasture component in silvopasture systems, increasing the productive and qualitative potential of forage from the herbaceous layer (Figueiredo et al. 2023). Legume trees can provide an important ecosystem service through the incorporation of litter high low C/N ratio increasing mineralization rates and nutrient cycling (Lira Jr. et al., 2020; Moreno-Gálvan et al. 2023); and improving the quality of soil through the storage of soil organic carbon (López-Hernandez et al. 2023).
The greater hemicellulose in plants during the dry period is a response to adverse environmental conditions. During periods of drought, plants are exposed to elevated solar radiation, higher temperatures, and less rainfall. Plants subjected to these extreme conditions can adopt anatomical and cellular modifications, including thicker cell walls and a greater leaf proportion of sclerenchyma and vascular tissues (Kering et al. 2011; Habermann et al. 2021). The increase in hemicellulose in the plant can bring some benefits to grazing animals, due to the ease for microorganisms to degrade this fraction in the digestive tract of ruminant animals compared to cellulose and lignin (Li, 2021). Increased ADF and NDF levels reduce fiber digestion, limiting forage consumption by ruminants and consequently can limit animal performance (Habermann et al. 2021).
Lignin synthesis requires high energy and nutrient uptake, which can be limited during drought stress (Peracchi et al. 2024). Furthermore, this fraction is richer in carbon and less hydrated when compared to hemicellulose and cellulose fractions, which can increase the demand for water for the synthesis of this compound (Rupitak and Srisaikam, 2021). In addition, the lignin content may vary over time during plant growth. M. caesalpiniifolia likely accumulated more hemicellulose in the dry period than lignin (Fig. 7A), considering the water demand and energy expenditure to synthesize this fibrous fraction, this assumption can be confirmed with the reduction in the content of ADF and lignin in the second year in the silvopasture (Fig. 7B).
The considerable CP content and the reduced NDF (Table 4), ADF, and lignin (Fig. 7B) in the second year contributed to increasing the digestibility of forage of M. caesalpiniifolia during the rainy season. Seasonal fluctuations occur throughout the year (Fig. 2), in the dry period, there was a reduction in forage quality, characterized by an increase in fiber content, a reduction in protein levels and digestibility (Fig. 8). Favorable climatic conditions with adequate rainfall and moderate temperatures contribute to greater forage quality and nutritional value (Tlahig et al. 2024).
In the nutritional aspect, IVDMD is also directly related to the ruminal concentration of volatile fatty acids (VFAs), which depends on the level of food intake and diet composition (Hall et al. 2015). Santos et al. (2020) evaluated in vitro ruminal fermentation parameters and methane production in Urochloa brizantha in a silvopasture system associated with levels of protein supplementation and reported that when there was a greater amount of NDF there was lower digestibility of the forage. When forage had a higher concentration of CP resulted in greater digestibility and, consequently, higher concentrations of VFAs. Forage digestibility is one of the essential measurements that can reveal the potential to provide energy for ruminants (Van Soest, 1994). Therefore, understanding variations in the nutritional value of the forage over time is an important advance to assist in the management of the arboreal component in silvopasture systems (Chebli et al. 2022; Santana et al. 2022). In the present study, IVDMD can be considered low, as it is outside the range (600–670 g kg− 1) as reported by Lascano (1994) for in vitro assays.
Another factor that must be taken into account in the nutritional value of forage from legume trees is the presence of secondary compounds, such as tannin, which can affect the acceptability of the forage by the grazing animals, resulting in a reduction in voluntary intake and dry matter digestibility (Guimarães-Beelen et al. 2006). Diniz et al. (2024) evaluated the nutritional value of Stylosanthes spp. a subshrub legume, and reported that the highest concentration of condensed tannins occurred in the dry period since the biosynthesis of condensed tannins is related to environmental conditions. Also, other factors such as the age of the plant and other abiotic factors associated with temperature changes, water content in plant tissues, photosynthetic radiation, and nutrient deficiency can modify the percentage of secondary compounds in legume plants (Oliveira et al. 2022). Condensed tannins represent substances that can negatively affect the use of nutrients by ruminants, due to their anti-nutritional characteristics (Costa et al. 2021).