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  • Review
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3 August 2026

18 Pages

Adaptation of Chloris gayana to Abiotic Stress Conditions: A Review

and
1
School of Agronomy, University of Buenos Aires, Buenos Aires C1417DSE, Argentina
2
Instituto de Investigaciones en Biociencias Agrícolas y Ambientales (INBA), Facultad de Agronomía, Universidad de Buenos Aires, Buenos Aires C1417DSE, Argentina
3
Academia Nacional de Agronomía y Veterinaria, Buenos Aires C1417DSE, Argentina
*
Author to whom correspondence should be addressed.

Abstract

Chloris gayana Kunth is a cultivated forage with worldwide distribution, based on its wide environmental adaptability. This review addresses some of the recent findings on C. gayana abiotic stress tolerance resulting from field or controlled conditions trials, focusing on salinity and alkalinity, drought, heat, and water excess. Management of the species under restraining environments is considered—a topic that most published information has concentrated on since a comprehensive review that was published in 2004. The main novel findings since then refer to the growing relevance role C. gayana is adopting in saline and alkaline soil amelioration, either through its association with mycorrhizal fungi or with trees in silvopastoral systems. This key trend is highlighted as a promising area for the development of sustainable production practices. Research gaps are identified. Among them, the development of standardized methodology guidelines for cultivar evaluation to guide adoption decisions and breeding efforts to improve its nutritive value under stress conditions. Tools for molecular intervention of the species that have become available since the previous reviews are also mentioned.

1. Introduction

Chloris gayana Kunth (Poaceae, a member of the subfamily Chloridoideae) is among the cultivated forages that sustain livestock production worldwide. It is a perennial stoloniferous grass native to Africa, introduced into cultivation in S. Africa in 1885 by Cecil Rhodes (hence its common name, Rhodes grass). From the early years of the 20th century, Rhodes grass expanded as a forage species throughout tropical and subtropical regions of the world. Its worldwide distribution is based on its broad temperature adaptability (from 35 °C to subzero), moderate rainfall requirements, and capacity to grow and thrive in a wide variety of soils.
Several botanical features have contributed to C. gayana’s extensive adoption as a forage: perennial, tufted plants with erect or ascending stems and abundant stolon production that provide rapid and efficient soil cover, wide soil and environmental adaptability [1,2]. It is productive for about 5 years, and it can withstand trampling [3].
C. gayana is an outbreeding (cross-pollinating) species featuring rich intravarietal variability. Such intravarietal variability has been confirmed in molecular genetic studies [4,5,6,7], and has been the basis for breeding cultivars which are essentially stabilized populations of either diploid (n = 20) or tetraploid (n = 40) specimens [8]. Diploid cultivars tend to flower earlier, exhibiting lower quality in the summer than tetraploid cultivars. In general, flowering in tetraploids responds to decreasing photoperiod, at the end of summer, which allows for the grazing period to be extended while maintaining good nutritional value during spring and summer. Diploids are more stoloniferous, while tetraploids are more erect and have more tillers. Cultivars have been obtained in countries as diverse as Japan, Australia, the USA, Kenya, Argentina, Zimbabwe, Tanzania, Uganda, Zambia, etc.
In addition to being used as forage, C. gayana supplies a range of ecosystem services, such as contributing to soil cover and to manage soil hydrology. It is also noted for its ability to stabilize disturbed sites [2], being effective for land reclamation and soil amelioration.
Context: Rhodes grass rapid expansion, broad environmental adaptability and multiple uses initially stimulated intensive botanical and physiological studies in the 20th century. This species was the subject of some of the earliest descriptions of C4 photosynthesis [9]. Comprehensive reviews on this species were written in 1969 by Bogdan [1], and by Loch et al. [2] in 2004. Those reviews also provide detailed botanical descriptions of the species and its environmental responses, as well as cultivar characteristics.
Justification: This review will mostly update research on abiotic stress effects on C. gayana published after the review by Loch et al. [2]. However, saline stress has been most intensively studied, followed by drought, while heat, cold, nutrient stress, and multi-stress performance remain under-characterized [4,10,11].
Objective: The review will focus on salinity, drought and heat, the main abiotic stress conditions that prevail in the areas where C. gayana is grown, and briefly mention other less generalized stresses.
Nowadays, abiotic stresses are usually inscribed within the concept of climate change conditions. These include increasing temperature, changing precipitation pattern, increased concentration of CO2 and ozone, and sudden and erratic occurrence of extreme meteorological events [12]. Global warming is expected to increase the frequency and intensity of droughts worldwide [13]. This combination of factors can also impose changes in soil salinity conditions (mainly increases) and desertification. Climate changes have stimulated the introduction of high-yielding subtropical grasses into temperate grasslands to increase forage offer [14], and C. gayana has been mentioned as one of the most promising species in this new scenario. While it is obvious that abiotic and biotic stresses co-exist and that plants are not exposed to individual stresses in the field but rather to a combination of stress conditions, the topical division in this article is meant only to provide an outline for examining some of the published research. The paucity of multi-stress studies is highlighted.
While C. gayana is considered as a broadly stress-adapted forage species, its agronomical performance depends strongly on cultivar characteristics, stress type, and growth stage. Some field management practices to alleviate stresses, and the effects of this species on soils will also be briefly considered; however, cultivar characteristics and adaptability will be mentioned succinctly as they have been previously reviewed [1,2], and many of the recent studies mention only the species, but omit specifying the cultivar used.

2. Tolerance to Saline, Alkaline and Saline–Alkaline Soil Conditions

Salinity is one of the most widespread soil constraints to agriculture across a wide range of climatic regions [15]. Salt-affected soils are a group including saline, saline–alkaline and alkaline soils which have a common origin and some similar properties. Saline soils have high concentrations of soluble salts, mainly chlorides and sulphates, while alkaline soils are characterized by high levels of exchangeable sodium. Transitions among types of soils within this group are common [15,16].

2.1. Salinity Tolerance

Salinity imposes both water-stress and ion-specific limitations in plants, which, in turn, may result in ion imbalance and toxicity [15,17]. Salt tolerance is classically defined as the ability of plants to maintain growth and productivity under saline conditions [18]. Species within the Poaceae family show a very wide range of salinity tolerance [19]. Among other stresses, salinity studies are the most numerous in C. gayana and it is recognized as one of the most salt-tolerant C4 forage grasses [2].
In agreement, Salinity Tolerance Index (STI) values characterized by Tobar Gómez et al. [20] have identified C. gayana as a salt-tolerant species. Table 1 summarizes two experiments evaluating the effect of salinity on C. gayana biomass production. The electrical conductivity (EC) in the substrate was 0, 4 to 5, and 8 to 9 dS m−1. Shoot biomass decreased significantly in both experiments at the highest salinity level. Root biomass decreased only in one experiment. These results are consistent with previously reported trials, which indicated that biomass production is affected when salinity exceeded EC 8 dS m−1 [2,3]. However, this salt concentration is far from the salt tolerance limit for this species and indicates that this grass can be grown in areas where other species cannot be cultivated [15,16].
Table 1. Shoot and root dry biomass (g/pot) from two pot experiments with C. gayana subject to two levels of salinity and a non-salinized control. Results are means ± standard deviations of six replicates per treatment. Data taken with permission from [20].
Diploid cultivars have been suggested to be more salt-tolerant than tetraploids [4]. That study indicated that tetraploid cultivar Boma was relatively less salt-tolerant than the diploids, while no differences in salt sensitivity were found among diploid cultivars surveyed in germination, early vegetative growth, ion accumulation or osmotic adjustment. In the United Arab Emirates, Abu-Alrub et al. [21], studied the performance of C. gayana cv. Katambora in the field, under controlled levels of soil salinities over two consecutive years. They evaluated potential yield and nutritional values in response to four saline irrigation levels (200, 6000, 12,000, 18,000 ppm). The highest dry matter yield was observed under 6000 ppm NaCl, showing, however, significant yield reductions at 12,000 ppm. Collectively, these examples highlight that while C. gayana can be considered a salt-tolerant species, in general terms, reductions in biomass accumulation can be expected at high salinity levels.
Salt-tolerance mechanisms have been extensively reviewed in the literature; [17,22,23,24,25] are some of the most recent ones, but there are many more. Illustrative diagrams and tables including the underlying molecular components to salt tolerance mechanisms can be consulted there. Briefly, salinity imposes water and ion limitations on plants. These can cause nutritional imbalances and toxicity, as well as alteration in reactive oxygen species (ROS) homeostasis and redox regulation. Initially, plants respond mainly to low water potentials that characterize saline substrates, while ion-specific effects are observed later, when potentially toxic ions that are in excess in saline substrates, such as Na+ and Cl−, accumulate internally. In turn, accumulation of essential elements, for instance K+, Ca2+, and N may be altered under these conditions. Water imbalance within cells may also result from apoplastic ion excess. Therefore, as summarized by Taleisnik et al. [18], adaptations to salt stress include: (a) water balance rendered by the accumulation of organic solutes in the cytosol and ions in vacuoles, (b) access restriction of potentially toxic ions to the sites of active metabolism, (c) the maintenance of intracellular concentrations of essential nutrients required for normal metabolism, and (d) detoxification of ROS. Three groups of mechanisms contribute to plant salinity tolerance, which are not exclusive and can operate either simultaneously or not. The first includes features that confer tolerance to substrate low water potential caused by salinity, through the accumulation of organic and inorganic solutes that lower internal water potential and generate the necessary gradient to ensure plant water uptake. The second participates in the control of the uptake and internal distribution (exclusion from sites of active metabolism) of potentially toxic ions, and the maintenance of intracellular concentrations of essential nutrients. The third type of salinity tolerance mechanisms provides tolerance to toxic ions in leaf tissues and also includes those involved in the control of oxidative stress. The antioxidant defense system includes enzymes such as superoxide dismutase, catalase, and peroxidase, as well as non-enzymatic antioxidants such as ascorbic acid, glutathione, and others. These antioxidants control lipid peroxidation, protein oxidation, and DNA damage caused by ROS, thereby stabilizing redox balance, maintaining cellular homeostasis, and improving plant tolerance to stress [26]. Malondialdehyde concentration, a marker for lipid peroxidation and thus, oxidative stress, was suggested as a useful tool to select salt-tolerant C. gayana individuals at initial growth stages [27]. As expected, all salt-tolerance mechanisms are triggered and coordinated by complex networks of signaling and gene expression regulation as detailed in the aforementioned reviews. Some components of the three types of salt-tolerance mechanisms have been identified in C. gayana [18]: ion secretion from salt glands (detailed below), antioxidant enzymatic system stimulation, and osmolyte accumulation. However, the underlying molecular aspects have not been explored, though they may be expected to be similar to those identified in other species.
In the Chloridoiedeae, salt tolerance has been associated, among other traits, with the exclusion of excess toxic ions, and, in some cases with ion secretion from leaf salt glands [19,28]. The discovery of bicellular salt glands on leaves of this species [29] was conceptually associated with its documented salt tolerance [1]. Salt secretion (also referred to as recretion or excretion) through salt glands plays a significant role in reducing ion concentration in shoots [30]. Zhang and Shi [31] describe salt gland-mediated ion exclusion as a hallmark of halophytic grasses. Several recent reviews deal with these structures [30,31,32,33]. de Luca et al. [34] and Kobayashi et al. [35] showed that these glands secrete both Na+ and K+, with a greater capacity for Na+ secretion. This reduces intracellular sodium accumulation and mitigates ionic toxicity. Additional cellular mechanisms, including Na+/H+ antiporters involved in vacuolar compartmentalization and pH regulation, likely support ionic homeostasis.
As mentioned earlier, diploid C. gayana cultivars are considered to be more salt-tolerant than tetraploids [4]. Some of the most conspicuous physiological differences between tetraploid cv. Boma and diploid cultivars in the response to salinity were the relatively lower Na+ and higher K+ accumulation in leaf tissues, and the lower Na+ excretion via salt glands in Boma [36]. Salt glands are easily distinguishable on leaf surfaces and provide a useful selection trait. Salt gland density was significantly higher in a salt-tolerant than in a salt-sensitive clone of C. gayana cv. Boma [34]. This clone was one of the plants used to develop the first registered tetraploid salt-tolerant cultivar, C. gayana cv. Epica INTA-Peman [37]. Salt gland density was one of the criteria used to develop salt-tolerant diploid cultivars from Topcut and Finecut [38].
A reduction in toxic ions (Na+ and Cl−) accumulation contributes to buffering the negative effects of salinity on metabolism. The distribution of Na+ and Cl− in leaf tissues of C. gayana plants grown under salinity was investigated by Oi et al. [39]. They found that Na+ and Cl− accumulated to the highest levels in xylem parenchyma and epidermal cells, but were maintained at lower concentrations in photosynthetically active mesophyll and bundle sheath cells, and lower in chloroplasts of those cells than in their vacuoles. Though the mechanisms responsible for such distribution were not explored, they concluded that it contributed to the lack of negative effects of salinity on photosynthesis.

2.2. Tolerance to Alkaline and Saline–Alkaline Soil Conditions

Alkaline soils show high pH (often >8.5), high HCO3− and CO32− concentrations, and are characterized by high exchangeable sodium percentage (ESP), reduced nutrient availability (specifically Fe and Zn), impaired soil structure resulting in limited water and air mobility and storage, intensive colloid eluviation, and dispersion of organic matter. Saline–sodic soils combine characteristics of both saline and alkaline soils, although salinity predominates; however, alkaline stress appears more damaging than salinity alone. Therefore, agricultural productivity of alkaline and saline–alkaline soils is restricted by their agrophysical, physiological and hydrological properties [16,40].
Some reports indicate that C. gayana is capable of tolerating elevated bicarbonate concentrations and high pH during germination and early establishment [41,42]. Kumar [43] observed that while early growth in alkali soils was reduced, once established, it achieved growth rates comparable to those in non-alkaline soils. Alkaline stress can inhibit root apical growth, affecting the emergence of root hairs and, subsequently, water and nutrient uptake. Alkalinity also affects chlorophyll content and may cause a significant accumulation of ROS. In turn, oxidative stress can also inhibit chlorophyll synthesis and accumulation. Saline–alkaline stress also decreases the K+/Na+, Ca2+/Na+, and Mg2+/Na+ ratios in shoots and roots [44].
Kumar [45] evaluated long-term forage yields on extremely sodic soil with an initial pH of 10.6 and ESP of 94. Under those conditions, Rhodes grass maintained higher productivity over time than other tropical grasses tested (Leptochloa fusca, Brachiaria mutica, Cynodon dactilon and Panicun antidotale) and was thus recommended for extremely sodic soils. Nevertheless, other studies indicate a reduction in C. gayana productivity with increasing pH or under combined salinity–alkalinity stress [46,47].
Table 2 summarizes information on biomass production of a C. gayana stand in a very alkaline soil, in comparison with Panicum coloratum L. and native vegetation [48]. The average annual forage production of the two forage grasses was similar (ranging from 4 to 7.5 × 103 kg/ha dry matter per year, depending on the rainfall), and higher than native vegetation. Clearly, the incorporation of both species brought about a marked increase in biomass production compared to the productivity of native vegetation. However, given their similar productivity, these results indicate that the selection of one grass over another for cultivation in a specific site depends on several factors. These include, among others, seed availability, tolerance to other stresses, the time of year when peak biomass production occurs, tolerance to livestock trampling, etc.
Table 2. Chloris gayana and Panicum coloratum annual shoot dry biomass production (kg/ha) during 5 years on an alkaline soil of the Flooding Pampas (Argentina). Means followed by different letters are significantly different (p < 0.05). Data taken with permission from [48].
The comprehensive review by Cao et al. [49] highlights the complexity of alkaline stress and summarizes morphological and physiological adaptations of plants growing in saline–alkaline soils and addresses the molecular networks underlying these responses. A section devoted to alkaline stress in Liang et al.’s review [50] focuses on molecular signaling elements. Analogous mechanisms are likely to be present in C. gayana; however, we have not been able to detect specific information for this species. As mentioned earlier, oxygen supply limitations and nutrient deficiencies contribute to growth restrictions in alkaline soils [51,52]. Experiments in P. coloratum distinguished some of the physiological causes for reduced growth under alkaline conditions [51]. Unfortunately, no such experiments have been carried out in C. gayana, and they could render useful information to orient breeding efforts.
In general, studies focusing on C. gayana under alkaline stress are sparse and usually include only one cultivar or compare species rather than cultivars [46,53]. As alkaline soils occupy some of the climatic regions where C. gayana thrives, specific studies on alkaline stress tolerance in this species, with emphasis on cultivar performance, would provide a scientific basis for breeding adequate forage grasses for such areas.

3. Temperature and Drought Adaptation

As mentioned in the Introduction, climate change predicts the occurrence of increasing temperature and concentration of CO2, changing precipitation patterns and erratic occurrence of extreme meteorological events [12]. These conditions, which obviously affect normal plant physiology, have not been extensively explored in C. gayana recent research. Nonetheless, general trends can be derived from studies on other C4 grasses.
For each plant species, a defined range of maximum and minimum temperatures forms the boundaries of observable growth [54]. Cardinal temperatures are the minimum (Tb), optimal (To), and maximum (Tm) for a specific developmental stage. Vegetative development rates gradually increase as temperatures rise to the species optimum level, and optimum temperatures are usually higher than for reproductive development. Cardinal temperatures for sowing to plant emergence were 13, 31 and 47 °C for Tb, To, and Tm, respectively [55]. Other studies indicate a maximum Tm of 50 °C [56].
Loch et al. [2] state that, in agreement with what is known for other C4 species, Rhodes grass grows best at high temperatures, an optimum of about 35 °C for photosynthesis was reported in growth chamber studies. Bogdan [1], citing evidence from trials in several countries, also indicates that the species is adapted to high temperatures, yet it can eventually withstand sub-zero temperatures. In Argentina, the species has been sown in areas prone to sub-zero temperatures in winter [57]. Nevertheless, in general, C. gayana is sensitive to frosty winters that can lead to plant death, as Wu et al. [58] report for Southern NSW (Australia).
Negative effects of heat stress on plants are well known, affecting both the light (electron transport) and dark (Calvin cycle) reactions of photosynthesis and rendering this process among the most thermosensitive aspects of plant physiology. In C4 species, elevated CO2 frequently decreases photosynthetic thermotolerance [59].
On the other hand, C4 species have greater rates of CO2 assimilation than C3 species for a given level of leaf nitrogen concentration, when both parameters are expressed either on a mass or an area basis. The lower nitrogen requirement of C4 plants results from their CO2-concentrating mechanism, which raises the bundle sheath CO2 concentration, saturating RuBisCo in normal air and almost eliminating photorespiration (RuBisCo is Ribulose-1,5-bisphosphate carboxylase/oxygenase; the enzyme has a dual role of catalyzing CO2 fixation and oxygenation of ribulose-1,5-bisphosphate, the second reaction initiates the process known as photorespiration). In turn, the mentioned CO2-concentrating mechanism is responsible, along with lower stomatal conductance, for the higher leaf and plant water-use efficiency of C4 plants. These characteristics are fully realized at high light and temperature [59]. Such mechanisms have led physiologists to conclude that C4 grasses are predicted to thrive under climate change in high temperature and drought, yet underscore that few studies have examined how these species will respond to combined stresses [60].
Rhodes grass is comparatively more drought-tolerant than other warm-season forage crops [61]. Loch et al. [2] cite water requirements to be in the range of 600 to 1200 mm, and report that Callide, a major tetraploid cultivar, has extended into wetter districts (to 1500 mm average rainfall), while Bogdan [1] reviewed evidence that C. gayana requires irrigation to grow in areas that receive less than 70 mm rainfall a year.
High temperatures and water stress often coincide. As expected, field research in low-desert and semi-arid environments consistently describes Rhodes grass as tolerant of high temperatures. Bachie [62] reported that, in desert conditions in California, C. gayana demonstrated strong adaptability and high biomass productivity under extreme summer heat. In agreement, Mganga et al. [63], working in semi-arid African rangelands under natural rainfed conditions and high ambient temperatures, documented that this grass-maintained productivity in landscapes exposed to intense seasonal heat. Studies in Brazil’s semi-arid region also support heat adaptation. Nobre et al. [64] found that genotypes such as Callide, Mariner, and Sabre performed well in regions characterized by low rainfall and high temperatures, implying cultivar-level adaptation to heat stress. The three cultivars are indicated as suitable genotypes for pasture establishment in the northern region of Minas Gerais, Brazil.
Several studies support the notion that drought tolerance and variability characterize this species, among them [64,65,66,67,68]. Trials under controlled or semi-controlled conditions show survival under prolonged drought. Wehr et al. [42] demonstrated that C. gayana survived up to 88 days without water and resprouted after six months of dry conditions. Chiacchiera et al. [69] showed that individual plants survived extended drought periods even when soil water potential dropped below the permanent wilting point. Field trials with 62 accessions from the International Livestock Research Institute Genebank in Ethiopia [65] revealed that while overall productivity declined under water deficit, several accessions maintained comparatively high biomass yields, suggesting both drought resilience and significant genotype-dependent variation. Brunetti et al. [70] simulated forage growth in Brazil under climate change and found increased drought stress during winter and spring, and intensified seasonality, which led to a slight decline in forage yield. Nevertheless, they expect that drought stress magnitude and period will be more predictable in the future and that some forages, such as the tropical Urochloa brizantha and Megathyrsus maximus, can contribute to reversing the trend in grass yield decline.
In sum, it can be said that C. gayana is moderately drought-tolerant; variability for this trait is found within both diploid and tetraploid germplasm of the species. This variability has traditionally been the source of germplasm for breeding schemes, leading to new cultivars. In forage grass breeding programs, drought tolerance is defined by criteria of yield maintenance or/and/or high yield relative to other genotypes. Physiological and molecular traits involved in drought tolerance in Poaceae grasses have recently been reviewed by Pirnajmedin et al. [71]. The authors highlight that escape, avoidance, tolerance, and drought recovery mechanisms participate in the response to drought and describe the meaning of each one, indicating that they are not mutually exclusive and are often combined consecutively or simultaneously. As adaptation to drought is a complex combination of many traits, controlled in turn by multiple gene networks, breeding for a single trait will render only limited results.

4. Tolerance to Water Excess and Acidity

As stated earlier, climate change includes sudden and erratic occurrences of extreme meteorological events [12]—among them, water excess. Waterlogging effects strongly depend on the water quality and duration of the stress. C. gayana can tolerate temporary waterlogging, but plants are killed by deep or prolonged flooding [2]. C. gayana seed germination percentages at 75% and 125% field capacity were around 20% in both cases [3]. Seedling loss during establishment is associated with prolonged periods of waterlogging or flooding. Imaz et al. [72] showed that adult plants of cv. Finecut were able to tolerate temporary winter and spring flooding without exhibiting plant death. Tolerance to temporary waterlogging could be related to increased aerenchyma in roots, able to store oxygen transported from shoots, a trait that also supports tolerance in poorly drained soils [53]. Waterlogged plants can also tolerate subsequent drought [10].
Acid soils are not frequent in the environments where C. gayana thrives. Yet, some of its effects have been studied. Rhodes grass has some establishment problems on very acid soils, and it is stated that it is not very tolerant to acidity in later developmental stages [73]. However, in a greenhouse experiment, no significant differences were found in plant establishment and growth at various pH levels (3, 5, 7, and 9) [3]. Nevertheless, both establishment and biomass production showed somewhat higher values at pH 7, though below significance levels.

5. Crop Management Under Stress Conditions

Pasture establishment in environments prone to stress requires, in addition to adequate germplasm, suitable seeding techniques to ensure successful establishment. Once the pasture is established, defoliation strategies must be adjusted to produce high-quality forage and maintain pasture productivity and persistence [74].
Chloris gayana is appreciated for its ability to produce seeds, to cover ground, to tolerate clipping, grazing and trampling; successful establishments and high forage yields have been observed from the establishment year summer [65,75]. Germination and establishment are critical stages for pasture development. C. gayana stands are relatively easy to establish, and establish very fast in non-saline/alkali soils and adequate climates [76]. In normal soils, a good seedbed is usually prepared with proper tillage, and the seeds are sown broadcast or in rows. Direct seeding also applies. A roller is then used to compact the soil, as successful establishment depends strongly on achieving adequate contact between seeds and the soil. C. gayana is usually sown at a very shallow depth (1–2 cm), as seeds are very small and have limited reserves. Wu and Asaduzzaman [58] determined that seedling emergence mainly occurred from seeds located down to 2 cm depth, at 5 cm seedling emergence was very low, and at 10 cm depth no emergence occurred. Due to the reduced size of the seeds, they can be coated or mixed with a carrier [57,75]. Sowing density will depend on seed quality, and it is highly variable across different cropping areas, ranging from 1 to 8 kg/ha [76]. Sowing can take place in spring or early autumn, depending on the climatic characteristics of each location [57,75].
In saline soils, it is desirable to match sowing with the period of lowest possible surface salinity, which usually happens at the beginning of the rainy season. Sowing methods influence germination and establishment. A comparison between no-till and conventional sowing in saline and non-saline plots showed that no-till sowing was more effective in saline soil in terms of achieving germination, seedling survival and growth, while in non-saline soil, results did not differ between both sowing methods, as summarized by Pérez et al. [74], who also reviewed previous work on C. gayana by that research group. Additionally, sowing in saline–alkaline soils can improve soil properties [56,75,77], as mentioned below, in Section 6.
Germination rates in saline soils usually does not exceed 10%, and therefore high sowing densities are required or, since seeds may have dormancy, in some cases they are treated with hormones [57]. The goal is to achieve a stand with 20 to 30 plants per square meter.
Establishment on saline and alkaline soils is often erratic, uneven, and slow, and numerous cases of failed establishment have been recorded. In addition, the risk of surface encrusting is an additional problem. Increased rate of seedling mortality occurs in periods of high temperatures and droughts, leading to a reduction in the initial stand. Adequate radiation is also important for a successful establishment, but in South Africa it was found that excessive radiation can burn and kill seedlings [3]. In saline soils where seedling establishment is difficult, the ability to produce stolons that ensure ground cover renders effective advantages, as has been observed in Rhodes grass cv. Epica INTA Pemán. This is a tetraploid cultivar with enhanced salinity tolerance and profusely stoloniferous [37], allowing it to achieve rapid soil cover.
C. gayana seedlings compete poorly with weeds or natural vegetation during the post-establishment period, which strongly affects the species persistence. If the goal is to achieve a pure pasture, good control of existing vegetation must be exerted prior to sowing. Total (non-selective) herbicides can be used for effective management of weeds at the time of sowing, in combination with other control techniques [57]. Instead of sowing, in some countries, pasture establishment by rooted stolons is suggested [75].
Nevertheless, C. gayana can be cultivated in pure stands, intercropped with natural vegetation or with cultivated legumes [56]. In Ethiopia and East African countries, intercropping C. gayana and alfalfa (Medicago saliva) is the most successful perennial pasture, but it is also suitable to be grown in association with many other tropical legumes [56,78].
Native vegetation productivity in saline and alkaline soils is usually reduced, low quality and bare patches are often observed. In this context, the main objective of introducing C. gayana is to increase forage production compared to native vegetation. In addition, the species can fill a production gap that is generally summer-related or can be used as forage reserves by making hay bales and, less commonly, silage [56,74,75].
When germination is insufficient to achieve maximum ground cover, it is generally recommended to avoid grazing the plot during the establishment year. This allows for the development and rooting of stolons. Conversely, if significant ground cover has been achieved, the plot can begin to be lightly grazed during the first year [56].
Once the pasture is established, it is necessary to implement a defoliation strategy that allows for the production of forage of the required quality while maintaining productivity and persistence. C. gayana, as well as other perennial subtropical forages, has lower quality than temperate grasses; crude protein content is around 4–7%, and digestibility ranges from 50 to 80%. Drought, salinity, and waterlogging affect C. gayana quality in somewhat different ways: drought mainly lowers crude protein, affecting seasonal nutritive value, salinity affects yield rather than quality, and for waterlogging, evidence is scarce and indirect. On the other hand, while protein and digestibility can increase in individual plants under saline conditions, if yield decreases, total grass quality will decrease per surface unit [20]. Canopy management can influence cold tolerance in C. gayana. In higher latitudes, where plants would be exposed to cold stress, it is recommended to defer grazing to the fall, so as to increase the chances of winter survival and spring resprouting [72].
Attempts to estimate the effects of climate change on forage quality do not show conclusive results. While for C3 forage plants, a generalized reduction in N concentration is reported [79], this cannot be generalized for C4 plants [80]. These are characterized by high productivity and adaptability to warm and dry climates and by better water and nitrogen use efficiency than C3 crops. Due to the photosynthetic characteristics of C4 plants, their yields will not directly benefit from elevated CO2, as in C3 plants, but their nutritional value is not predicted to be negatively affected. Jobe et al. [81] outline possible research questions that will contribute information to this knowledge gap. To the best of our knowledge, no such studies have been conducted in C. gayana.
Proper forage quality for animal nutrition requires higher cellulose and hemicellulose and lower lignin content. One of the factors determining forage quality is the leaf-to-stem ratio, as leaves generally have lower lignin and higher cellulose and hemicellulose content, while stems contain a higher proportion of lignin. Forage quality (protein, fiber, and digestibility) is not a static property, it decreases with aging, mainly due to physical and chemical changes in the cell walls of the supporting tissues. However, it can be modified and improved by controlling the defoliation regime; this is so because leaf blade digestibility increases and fiber decreases with plant regrowth after grazing [82]. Maintaining a high tillering rate and a balance between the emergence and death of tillers is highly dependent on the defoliation regime adopted, as it determines the evolution of the leaf area index (LAI), a key factor in controlling both the emergence and death of tillers. A high tiller population density is associated with pasture persistence [74]. Frequent grazing generally improves forage quality but may limit total pasture production, while less frequent grazing may allow for greater total forage accumulation but of lower quality [73,83].
Nitrogen (N) is the most limiting nutrient affecting Gramineae pasture species productivity. A post emergence application, in the case of pure stands, stimulates rapid growth and application after each harvest or grazing period, and maintains productivity and quality [75]. Fertilization with phosphorus (P) has an impact at the seeding time and plays a critical role in improving grass productivity and protein contents [12,57,84]. In adult plants, P nutrition is associated with larger root systems under drought [61] and deeper and extensive roots, improving access to water under drought [85]. The most common fertilizers are urea for N and superphosphates for P. The doses, the machinery used, time and frequency of application, etc., vary across countries and production systems. Fertilization is site-specific but not universal. For example, N improved biomass and crude protein on sodic Argentine soils, but it showed no significant agronomic response in Ethiopia [86,87].
Cattle grazing can recycle, through excreta, 60 to 90% of ingested nutrients back to the soil, depending on the composition of the forage, type of animal, and time of grazing, but the nutrients are not uniformly distributed across the pasture, increasing the problem of soil spatial variability [47].
Meat and milk production on C. gayana pastures vary according to the stocking rates adapted to each agrosystem, which in turn, depend on characteristics such as climate, management (fertilization, irrigation, etc.), age and/or cattle weight, type of production, among others. The final yield as meat is highly variable, in Argentina, with a stocking rate of 2/3 heifers/ha, weight gains were in the order of 300 to 500 g/day [88], while in Australia, with a stocking rate up to 4 animals/ha, the weight gains were in the order of 0.500 to 2.3 g/day [56]. In East Africa, the weight gains were in the order of 500 g/day with a stocking rate up to 4 animals/ha [75]. It is concluded that recommended management technologies and cattle stocking rates must be determined locally [66].
Microorganisms to mitigate salt stress and increase forage yield appear as positive management tools. Yañez Yazile et al. [89], in a study from germination to early growth under controlled greenhouse conditions, found that some halotolerant bacteria can improve C. gayana ion balance, antioxidant activity, and photosynthetic efficiency. However, beneficial effects of microorganisms in some other studies under saline conditions are either absent, conditional, or inconclusive [20,90].

6. Chloris gayana for Saline and Alkaline Soil Amelioration

Sown pastures in saline soils usually increase biomass production and soil cover, often doubling the cover achieved by previous halophytic vegetation [77]. Plant biomass lowers soil temperature, reduces the rate of water evaporation, and thus decreases the risk of salinization. In addition, Rhodes grass can control soil erosion in bare patches of salt-affected soils due to its stoloniferous growth and extensive, vigorous roots [73]. Generally, increased soil cover with live and dead biomass has been indicated as the cause of soil organic carbon (SOC) increases. However, these increases in SOC have not always been observed when native vegetation is replaced [73]. Otondo et al. [77] found in alkaline soils that the introduction of C. gayana plots in native vegetation gradually increased net primary production up to 57–74% after 5 years, as compared to the original pasture. The soil, however, only showed slight decreases in topsoil pH (always higher than 9) and no significant change in salinity and SOC.
However, the soil showed a significantly better physical environment in bulk density, soil bearing capacity (at low soil water content), and soil structure. The last measurement is the index obtained from the change in the weighted mean diameter of the aggregates. Soil structural aggradation (as opposed to degradation) increases under a C. gayana pasture (Table 3).
Table 3. Relative changes in soil physical properties, bulk density, soil bearing capacity (at low soil water content), and soil structure (index obtained from the change in the weighted mean aggregate diameter) after 5 years of C. gayana establishment. Data taken with permission from [77].
Association with mycorrhizal fungi (MF) is a complementary approach not only to enhance productivity and stability in marginal forage systems, but also to recover saline–alkaline soils [91,92]. Recent research highlights the role of C. gayana in SOC stabilization within saline pastures. Banegas et al. [93] demonstrated that MF significantly contributes to SOC in saline pasture systems. It is through the production of glomalin-related soil proteins, which account for approximately 5–10% of SOC and 5–13% of soil N and act as a long-term carbon sink with turnover times of 6–42 years. In saline soils, glomalin accumulation enhanced carbon stocks under tropical perennial grasses and can improve soil aggregation and stabilize SOC under saline conditions.
Silvopastoral systems, integrating salt-tolerant trees with perennial forage grasses, offer a biologically driven strategy for sustaining livestock production in such environments. Dagar et al. [94] emphasize that grasses such as C. gayana are viable silvopastoral components for highly sodic soils, improving soil fertility and SOC sequestration under changing climate scenarios. Recent studies from South Asia [94] and the South American Dry Chaco [93] demonstrate that C. gayana-based pastures and silvopastoral systems significantly enhance SOC and total N, particularly through mineral-associated carbon pools. In the Indo-Gangetic Plains, Singh et al. [95] confirmed that C. gayana maintained productivity under tree canopies, suggesting compatibility with light-modified environments typical of silvopastoral systems. They reported that the Acacia nilotica + C. gayana silvopastoral system produced maximum understory biomass and showed strong soil ameliorative effects. Chhabra [96] further documented the suitability of Rhodes grass for alkali soil reclamation, highlighting its biomass potential and adaptability. A recent six-year study [93] in sodic soils of the Dry Chaco compared pure pasture and silvopastoral systems cultivated with C. gayana cv. Epica INTA-Pemán beneath Neltuma alba (syn. Prosopis alba). Their results indicate that both pure pasture and silvopastoral systems enhance soil fertility (SOC and total N) in sodic soils, yet tree spatial configuration significantly influences carbon distribution.
Collectively, the above studies highlight that beyond the recognized salt tolerance and biomass productivity of C. gayana, its contribution to SOC and total nitrogen accumulation through mineral-associated carbon fractions and mycorrhiza-mediated glomalin deposition are important components of biologically based restoration strategies.

7. Concluding Remarks, Research Gaps and Prospects

Chloris gayana is regarded as a stress-resilient forage, and research from the end of the 19th century to the mid-20th century dealt mainly with technological issues. Subsequently, it focused on physiological aspects, including the mechanisms of stress tolerance, mainly those related to salinity. Among them, salt secretion from salt glands was described and measured. This basic knowledge stimulated and boosted the genetic improvement of this species. In recent years, research has turned again to technological topics. However, as Loch et al. [2] mention, despite the importance of this species, after the 1970s the intensity of research and breeding efforts have not been maintained. This is still true. Considering this situation, several knowledge gaps that deserve consideration are highlighted in the following paragraphs.
Tolerance mechanisms to stresses other than salinity have been less explored, and cultivar differences have not been consistently tested. Successful introduction and survival of C. gayana in higher latitudes, as is currently happening, will require reliable information on cold tolerance and cultivar adaptation. Some of these issues were brought up above, in the respective sections.
Additionally, most evidence relies on laboratory and greenhouse trials, which may not translate directly to field conditions, where combined stresses prevail. This can be mentioned not only for the most common abiotic stresses C. gayana is exposed to, but also for nutritional stress. Nevertheless, it must be acknowledged that the gap between studies performed under controlled and field conditions is a common problem in other species as well, not only in Rhodes grass [97]. Many discrepancies found in the literature are probably due to methodological procedures rather than to biological contradictions.
Yet the biggest challenge remains in multi-stress environments, which require trials that exceed controlled conditions, in real fields, while monitoring environmental and biological variables, so that comparisons among trials can be implemented. A change in paradigm from individual stresses to multi-stress conditions is essential to deliver realistic solutions that imply not just genotype development but proven management packages.
The development of standardized methodology guidelines for trials is essential to ensure local research efforts will contribute to producing a comprehensive global picture. Existing reviews of regional information on C. gayana, such as that developed for Argentina [98], must surely exist for other regions, and they should be integrated when developing management recommendations.
Cultivar evaluation across diverse environments is required to guide sowing decisions. Plant establishment remains a challenge for several stress conditions, and more so for combined environmental stress. While establishment will depend also on seed quality, sowing methods, and climatic effects, unless this gap is approached with standardized protocols, no generalized recommendations will result.
Relatively lower forage quality of C4 species in comparison to temperate zone forages is a weakness, amplified by the negative effects that stress exerts on quality. Improving the nutritional quality of subtropical and tropical forage grasses is another challenge [12]. While general differences between C. gayana tetraploid and diploid cultivars exist in this respect, favoring the former, and the effects of clipping schemes on quality are largely known, forage quality remains a challenging issue. In the context of forage quality, available information points to site-specific responses to N that need to be considered at a global scale. Understanding N cycling and characteristics of this species is critical for estimating optimal N requirements [99]. Furthermore, Qaderi et al. [80] underscore that, in C4 plants, the number of research studies that combine at least three main features of climate change: elevated CO2 concentration, high temperature and water stress on N assimilation, is very limited. A similar reasoning applies as well to other nutrients such as P and K.
C. gayana is considered a species with relevant environmental roles, participating in saline and alkaline soil amelioration, either by itself, in consortium with trees and bushes in silvopastoral systems, or through its association with bacteria [89] or mycorrhizal fungi, or in endosymbiosis with beneficial microbes. This is a growing area of research that will no doubt produce tangible information on the sustainability of pasture systems [91]. A systemic approximation to plants in association with microorganisms, rather than to isolated plants, in the holobiont concept, is another change in paradigm that has gradually become established. Increasing attention is being devoted to plant–microbe beneficial associations that may mitigate stress effects, an approach that also includes biostimulants. Going from controlled to field trials is, in many cases, a pending gap.
In crops, breeding leads to improvement in traits relevant to their performance for agriculture and the target environment, which includes tolerance/resistance to biotic and abiotic factors [100]. However, in forage crops, breeding foci are directed to additional traits such as the pattern of seasonal dry-matter production, nutritive value to grazing animals, grass–legume compatibility, response to management, and persistence [99]. Those authors indicate that breeding for new pasture crops has stagnated in recent years. Achon et al. [101] underscore that integrating conventional breeding with cutting-edge molecular tools is necessary for developing climate-resilient, nutritionally enhanced tropical forages to support the long-term sustainability of grass-based livestock systems. A genetic linkage map has been produced for Rhodes grass [102] as well as an annotated draft of the whole genome sequence [103]. This information, along with the availability of transformation protocols [104], will facilitate genomic breeding approaches and genome editing on this species, to develop new high-quality forage resources for regions where high temperatures or soil constraints are likely to reduce pasture quality and productivity. Improving forage yield and quality will no doubt have positive impacts on livestock farming and all its related stakeholders.

Author Contributions

R.S.L. and E.T. contributed equally to this review. All authors have read and agreed to the published version of the manuscript.

Funding

This review was partially funded by Collaborative Project U.E. #101082089 “Raising the bio-based industrial feedstock capacity of Marginal Lands” 2023–2026.

Data Availability Statement

No new data were created or analyzed in this study. Data sharing is not applicable to this article.

Acknowledgments

ET dedicates this review to the memory of Oscar “Cachi” Melo, an enlightening colleague who introduced her to the notion that salt-tolerant forage grasses are essential for sustainable cattle production in subtropical, semi-arid regions of Argentina.

Conflicts of Interest

The authors declare no conflicts of interest.

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