Biomass components and water use efficiency in cactus pear under different irrigation systems and harvest frequencies

The objective was to evaluate the effect of different irrigation systems and harvest frequencies on the cultivation of cactus pear. The study was conducted in the semi-arid region, in the municipality of Russas, CE, Brazil. The experiment was a split plot randomized block design, with four replications. Plots were represented by irrigation systems (surface drip, microsprinkler, Micro Spray Jet, and conventional sprinkler) and subplots by harvest frequencies (6, 9, 12, and 18 months). Cactus pear biomass production characteristics were evaluated. Most of the productive characteristics evaluated were influenced by harvest frequency and irrigation system. The harvest frequency of 12 months combined with conventional sprinkler system resulted in a total dry biomass production of 16,400 kg ha−1. The conventional sprinkler system showed the highest number of second-order cladodes (4.9 cladodes plant−1). The conventional sprinkler and Micro Spray Jet systems showed the highest values for various structural characteristics. For water use efficiency (WUE), the highest value was found at the frequency of six months (8.46 kg ha−1 mm−1). Regarding the effects of irrigation system on WUE, the highest values were observed with the conventional sprinkler system (8.20 kg ha−1 mm−1). The harvest frequency of 12 months combined with the conventional sprinkler system presents better results in the evaluated productive characteristics, mainly the cactus pear production by area.(AU)

Impact of the female and hermaphrodite forms of Opuntia robusta on the plant defence hypothesis.

The optimal defence hypothesis predicts that increased plant defence capabilities, lower levels of damage, and lower investment in vegetative biomass will occur more frequently in sexual forms with higher resource-demanding tissue production and/or younger plant parts. We aimed to examine the effects of sexual form, cladode, and flower age on growth rate, herbivore damage, and 4-hydroxybenzoic acid (4-HBA), chlorogenic acid, and quercetin (QUE) concentrations in Opuntia robusta plants in central Mexico. Our findings demonstrated that hermaphrodite flowers showed faster growth and lesser damage than female flowers. The effect of cladode sexual forms on 4-HBA and QUE occurrence was consistent with the predictions of the optimal defence hypothesis. However, chlorogenic acid occurrences were not significantly affected by sexual forms. Old cladodes exhibited higher QUE and 4-HBA occurrences than young cladodes, and hermaphrodites exhibited higher 4-HBA concentrations than females. Resource allocation for reproduction and secondary metabolite production, and growth was higher and lower, respectively, in females, compared to hermaphrodites, indicating a trade-off between investment in reproduction, growth, and secondary metabolite production. Secondary metabolite concentrations in O. robusta plants were not negatively correlated with herbivore damage, and the two traits were not accurate predictors of plant reproductive output.

Pre-zygotic reproductive isolation between two synchronopatric Opuntia (Cactaceae) species in the Brazilian Chaco.

Opuntia (Cactaceae) is known for high rates of hybridization and ploidisation, resulting in the formation of new species. The occurrence of two sympatric and closely related species of Opuntia, O. elata and O. retrorsa, in Brazilian Chaco enabled us to test the hypothesis that pre-zygotic reproductive isolation mechanisms operate in both species. We monitored the flowering period, as well as floral biology, and compared the morphological variation of floral structures through measurements, performed intra- and interspecific cross-pollination tests, and recorded the guild of floral visitors and pollinators. Flowering was seasonal and highly synchronous. Floral biology exhibits similar strategies, and although floral morphology differs significantly in many of the compared structures, such morphological variation does not result in the selection of exclusive pollinators. Floral visitors and pollinators are oligolectic bees shared by both species. Opuntia elata and O. retrorsa are self-compatible. While interspecific cross-pollination (bidirectional) resulted in germination, the pollen tube did not penetrate the stigma. Opuntia elata and O. retrorsa are closely related; however, they are isolated and do not hybridise in Brazilian Chaco. We found that both have weak pre-pollination barriers, but that they are strongly isolated by pollen-pistil incompatibility, i.e. post-pollination barrier.

Comparative development of staminate and pistillate flowers in the dioecious cactus Opuntia robusta.

KEY MESSAGE: PCD role in unisexual flowers. The developmental processes underlying the transition from hermaphroditism to unisexuality are key to understanding variation and evolution of floral structure and function. A detailed examination of the cytological and histological patterns involved in pollen and ovule development of staminate and pistillate flowers in the dioecious Opuntia robusta was undertaken, and the potential involvement of programmed cell death in the abortion of the sex whorls was explored. Flowers initiated development as hermaphrodites and became functionally unisexual by anthesis. Female individuals have pistillate flowers with a conspicuous stigma, functional ovary, collapsed stamens and no pollen grains. Male individuals have staminate flowers, with large yellow anthers, abundant pollen grains, underdeveloped stigma, style and an ovary that rarely produced ovules. In pistillate flowers, anther abortion resulted from the premature degradation of the tapetum by PCD, followed by irregular deposition of callose wall around the microsporocytes, and finally by microspore degradation. In staminate flowers, the stigma could support pollen germination; however, the ovaries were reduced, with evidence of placental arrest and ovule abortion through PCD, when ovules were present. We demonstrate that PCD is recruited in both pistillate and staminate flower development; however, it occurs at different times of floral development. This study contributes to the understanding of the nature of the O. robusta breeding system and identifies developmental landmarks that contribute to sexual determination in Cactaceae.

Accumulation and export of nutrients in cactus pear cladodes (Opuntia ficus-indica) under different managements in the Brazilian Semiarid

The present study evaluated the accumulation and export of nutrients in cactus pear (Opuntia ficus-indica) cv. Gigante, fertilized with combinations of nitrogen (10, 70, 100, 130, and 190 kg ha−1 year−1 N) and phosphorus (10, 70, 100, 130, and 190 kg ha−1 year−1 P2O5) under annual and biennial harvests, in Quixadá and Tejuçuoca, using a split-plot randomized block design with four replications. In Quixadá, under annual and biennial harvests, the following orders of accumulation of macronutrients (in kg ha−1 year−1) were found, respectively: K (98.8) > Ca (87.2) > N (46.7) > Mg (26.8) > S (18.4) > P (2.04) and Ca (33.5) > K (31.1) > S (18.6) > N (12.9) > Mg (10.5) > P (0.81). In Tejuçuoca, under annual and biennial harvests, the orders of accumulation of macronutrients were, respectively: K (146.5) > Ca (204.6) > N (128.1) > Mg (75.8) > S (50.3) > P (3.7) and K (397.2) > N (191.3) > S (241.2) > Ca (167.8) > Mg (131.0) > P (14.1). The maintenance/production fertilization in cactus pear should be planned according to productive potential, fertilization and harvest managements, and cultivation region, based on nutritional requirement and considering the nutrient recovery efficiency.(AU)

Effects of catalase on chloroplast arrangement in Opuntia streptacantha chlorenchyma cells under salt stress.

In arid and semiarid regions, low precipitation rates lead to soil salinity problems, which may limit plant establishment, growth, and survival. Herein, we investigated the NaCl stress effect on chlorophyll fluorescence, photosynthetic-pigments, movement and chloroplasts ultrastructure in chlorenchyma cells of Opuntia streptacantha cladodes. Cladodes segments were exposed to salt stress at 0, 100, 200, and 300 mM NaCl for 8, 16, and 24 h. The results showed that salt stress reduced chlorophyll content, F v /F m , ΦPSII, and qP values. Under the highest salt stress treatments, the chloroplasts were densely clumped toward the cell center and thylakoid membranes were notably affected. We analyzed the effect of exogenous catalase in salt-stressed cladode segments during 8, 16, and 24 h. The catalase application to salt-stressed cladodes counteracted the NaCl adverse effects, increasing the chlorophyll fluorescence parameters, photosynthetic-pigments, and avoided chloroplast clustering. Our results indicate that salt stress triggered the chloroplast clumping and affected the photosynthesis in O. streptacantha chlorenchyma cells. The exogenous catalase reverted the H2O2 accumulation and clustering of chloroplast, which led to an improvement of the photosynthetic efficiency. These data suggest that H2O2 detoxification by catalase is important to protect the chloroplast, thus conserving the photosynthetic activity in O. streptacantha under stress.

Cactus stem (Opuntia ficus-indica Mill): anatomy, physiology and chemical composition with emphasis on its biofunctional properties.

Cactus stem (Opuntia ficus-indica Mill) is native to Mesoamerica and marketed in different forms such as fresh, frozen or pre-cooked. Worldwide, this vegetable is recognized for its pharmaceutical actions, including its antioxidant, diuretic, anticarcinogenic, anti-inflammatory, anti-diabetic, and anti-hypercholesterolemic properties, as well as their antiviral and antispermatogenic effects. However, not all of these properties have been associated with its chemical composition; therefore, this review aims to present and integrate information available on the physiology and anatomy of cactus stem and its chemical composition, focusing on some of the many factors that determine its biofunctionality. © 2017 Society of Chemical Industry.

Chemical and physical defense traits in two sexual forms of Opuntia robusta in Central Eastern Mexico.

Sexually dimorphic plants provide an excellent opportunity for examining the differences in the extent of their defense against herbivores because they exhibit sex-related differences in reproductive investment. Such differences enable comparison of the sex with high reproduction expenses with the sex that expends less. The more costly sex is usually also better defended against herbivores. Generally, females are considered more valuable than hermaphrodites in terms of fitness; however, hermaphrodites are more valuable if they can produce seed by autonomous selfing, provided that the inbreeding depression is low and pollen is limited. We studied a gynodioecious population of Opuntia robusta from Central-Eastern Mexico, which has been reported to be trioecious, dioecious, or hermaphrodite, and addressed the following questions: 1) Is the hermaphrodite's reproductive output higher than the female's, and are hermaphrodites thus better defended? 2) Are plant tissues differentially defended? 3) Do trade-offs exist among different physical defense traits? and 4) among physical and chemical defense traits? We found that 1) hermaphrodites had a higher seed output and more spines per areola than females and that their spines contained less moisture. Non-reproductive hermaphrodite cladodes contained more total phenolic compounds (TPCs) than female ones. In addition, 2) hermaphrodite reproductive cladodes bore more spines than female cladodes, and 3) and 4) we found a negative relationship between spine number per areola and areola number per cladode and a positive relationship between spine number per areola per plant and TPC concentration per plant. Non-reproductive hermaphrodite cladodes contained a higher concentration of TPCs than female cladodes, and parental cladodes contained fewer TPCs than both reproductive and empty cladodes.

The importance of pollinator generalization and abundance for the reproductive success of a generalist plant.

Previous studies have examined separately how pollinator generalization and abundance influence plant reproductive success, but none so far has evaluated simultaneously the relative importance of these pollinator attributes. Here we evaluated the extent to which pollinator generalization and abundance influence plant reproductive success per visit and at the population level on a generalist plant, Opuntia sulphurea (Cactaceae). We used field experiments and path analysis to evaluate whether the per-visit effect is determined by the pollinator's degree of generalization, and whether the population level effect (pollinator impact) is determined by the pollinator's degree of generalization and abundance. Based on the models we tested, we concluded that the per-visit effect of a pollinator on plant reproduction was not determined by the pollinators' degree of generalization, while the population-level impact of a pollinator on plant reproduction was mainly determined by the pollinators' degree of generalization. Thus, generalist pollinators have the greatest species impact on pollination and reproductive success of O. sulphurea. According to our analysis this greatest impact of generalist pollinators may be partly explained by pollinator abundance. However, as abundance does not suffice as an explanation of pollinator impact, we suggest that vagility, need for resource consumption, and energetic efficiency of generalist pollinators may also contribute to determine a pollinator's impact on plant reproduction.

Programmed cell death promotes male sterility in the functional dioecious Opuntia stenopetala (Cactaceae).

BACKGROUND AND AIMS: The sexual separation in dioecious species has interested biologists for decades; however, the cellular mechanism leading to unisexuality has been poorly understood. In this study, the cellular changes that lead to male sterility in the functionally dioecious cactus, Opuntia stenopetala, are described. METHODS: The spatial and temporal patterns of programmed cell death (PCD) were determined in the anthers of male and female flowers using scanning electron microscopy analysis and histological observations, focusing attention on the transition from bisexual to unisexual development. In addition, terminal deoxynucleotidyl transferase-mediated dUTP nick-end labelling assays were used as an indicator of DNA fragmentation to corroborate PCD. KEY RESULTS: PCD was detected in anthers of both female and male flowers, but their patterns differed in time and space. Functionally male individuals developed viable pollen, and normal development involved PCD on each layer of the anther wall, which occurred progressively from the inner (tapetum) to the outer layer (epidermis). Conversely, functional female individuals aborted anthers by premature and displaced PCD. In anthers of female flowers, the first signs of PCD, such as a nucleus with irregular shape, fragmented and condensed chromatin, high vacuolization and condensed cytoplasm, occurred at the microspore mother cell stage. Later these features were observed simultaneously in all anther wall layers, connective tissue and filament. Neither pollen formation nor anther dehiscence was detected in female flowers of O. stenopetala due to total anther disruption. CONCLUSIONS: Temporal and spatial changes in the patterns of PCD are responsible for male sterility of female flowers in O. stenopetala. Male fertility requires the co-ordination of different events, which, when altered, can lead to male sterility and to functionally unisexual individuals. PCD could be a widespread mechanism in the determination of functionally dioecious species.

Effect of fungi and light on seed germination of three Opuntia species from semiarid lands of central Mexico.

Fungal attack under light reduces mechanical resistance of the testa of Opuntia seeds, making it easier for the embryo to emerge. However, the effect of fungi on Opuntia seed germination in darkness is unknown. We evaluated the combined effects of light and inoculation with Phoma medicaginis, Trichoderma harzianum, Trichoderma koningii, and Penicillium chrysogenum on germination of O. streptacantha, O. leucotricha, and O. robusta seeds, from central Mexico. We also evaluated the combined effects of seed age (2-, 3-, and 12-year-old seeds) and presence of fungi on the testa on O. streptacantha germination. All fungal species eroded the funicular envelope and promoted seed germination for O. leucotricha and O. streptacantha, but did more so in light than in darkness. For the latter species, younger seeds inoculated with fungi had lower germination than older ones. For O. robusta, we found that seeds inoculated with P. medicaginis and T. harzianum had similar germination in light and in darkness. Our results strongly indicate that deterioration of the testa by fungi is higher in light than in darkness.

Functional characterization of an acidic SK(3) dehydrin isolated from an Opuntia streptacantha cDNA library.

Cactus pears are succulent plants of the Cactaceae family adapted to extremely arid, hot and cold environments, making them excellent models for the study of molecular mechanisms underlying abiotic stress tolerance. Herein, we report a directional cDNA library from 12-month-old cladodes of Opuntia streptacantha plants subjected to abiotic stresses. A total of 442 clones were sequenced, representing 329 cactus pear unigenes, classified into eleven functional categories. The most abundant EST (unigen 33) was characterized under abiotic stress. This cDNA of 905 bp encodes a SK(3)-type acidic dehydrin of 248 amino acids. The OpsDHN1 gene contains an intron inserted within the sequence encoding the S-motif. qRT-PCR analysis shows that the OpsDHN1 transcript is specifically accumulated in response to cold stress, and induced by abscisic acid. Over-expression of the OpsDHN1 gene in Arabidopsis thaliana leads to enhanced tolerance to freezing treatment, suggesting that OpsDHN1 participates in freezing stress responsiveness. Generation of the first EST collection for the characterization of cactus pear genes constitutes a useful platform for the understanding of molecular mechanisms of stress tolerance in Opuntia and other CAM plants.

Drought-stress-induced up-regulation of CAM in seedlings of a tropical cactus, Opuntia elatior, operating predominantly in the C3 mode.

Immediately after unfolding, cotyledons of the tropical platyopuntoid cactus, Opuntia elatior Mill., exhibited a C(3)-type diel CO(2) exchange pattern characterized by net CO(2) uptake in the light. Significant nocturnal increases in titratable acidity typical of crassulacean acid metabolism (CAM) were not detected at this early developmental stage. As cotyledons matured and the first cladode (flattened stem) developed, features of CAM were observed and the magnitude of CAM increased. Nonetheless, in well-watered seedlings up to 10 cm tall, C(3) photosynthetic CO(2) fixation in the light remained the major pathway of carbon fixation. Reduced soil water availability led to an up-regulation of net dark CO(2) fixation and greater nocturnal increases in tissue acidity, consistent with facultative CAM. These observations demonstrate that C(3) photosynthesis, drought-stress-related facultative CAM, and developmentally controlled constitutive CAM can all contribute to the early growth of O. elatior. The strong C(3) component and facultative CAM features expressed in young O. elatior contrast with mature plants in which obligate CAM is the major pathway of carbon acquisition.

Prickly coexistence or blunt competition? Opuntia refugia in an invaded rodent community.

Endemic Nesoryzomys swarthi and invasive Rattus rattus exist in unlikely sympatry in Galápagos as female N. swarthi suffer from competition with R. rattus. This study evaluates the role of feeding habits in facilitating their co-occurrence. Spool-and-line tracking of 85 N. swarthi and 33 R. rattus was used to quantify their selected diets, foods of which were used in captive trials of 46 N. swarthi and 34 R. rattus to quantify their preferred diets. Selected diets were compared between species and seasons using niche measures, and contrasted to preferred diets to qualify inferences about competition. Diet overlap was highest in the wet season when food-particularly fruit-abundance was highest and R. rattus diet breadth was broadest. Preferred and selected diets were marginally correlated for R. rattus but uncorrelated for N. swarthi, suggesting that R. rattus interfere with N. swarthi foraging. Diet overlap was highest between female N. swarthi and R. rattus, perhaps due to female breeding requirements. Male N. swarthi avoided R. rattus-preferred foods, possibly to avoid aggressive encounters with R. rattus. During the dry season, when foods declined and the R. rattus population crashed, diet overlap was lowest as R. rattus diet narrowed in the absence of fruits. Female, and particularly male, N. swarthi diet broadened, with emphasis on acquiring Opuntia foods but N. swarthi-preferred and selected diets were uncorrelated, suggesting that their foraging was inhibited by R. rattus. In conclusion, the narrower diet breadth of R. rattus in contrast to N. swarthi suggests that this species may be less adapted to food acquisition at this site, particularly when fruits are absent. The year-round presence of R. rattus, however, appears to inhibit N. swarthi foraging for its preferred diet, and they instead specialise on Opuntia foods, which were uneaten by R. rattus and may provide N. swarthi with a localised competition refuge from encounters with R. rattus.

Biochemical and nutritional characterization of three prickly pear species with different ripening behavior.

Biochemical and nutritional changes were studied during the ripening process of three Opuntia morphospecies with different ripening behavior: Naranjona (O. ficus-indica), Blanca Cristalina (Opuntia sp.), and Esmeralda (Opuntia sp.) of early, early-intermediate, and intermediate-late ripening, respectively. In loss of fresh weight, Naranjona showed the highest values, while in Blanca Cristalina and Esmeralda, a discrete weight loss was found. No significant differences were found among morphospecies in soluble solids, total titratable acidity and pH during the postharvest days. Blanca Cristalina and Esmeralda showed an increase in the content of carotenoids, while these diminished in Naranjona. The cell wall enzymes evaluated showed particular behaviors during the ripening of each morphospecies suggesting a fine biochemical control and not a clear relationship between fruit softening and enzyme activity. This study provides basic information on prickly pear ripening, in order to understand this process for its control and for improving shelf life.

Young daughter cladodes affect CO2 uptake by mother cladodes of Opuntia ficus-indica.

BACKGROUND AND AIMS: Drought damages cultivated C3, C4 and CAM plants in the semi-arid lands of central Mexico. Drought damage to Opuntia is common when mother cladodes, planted during the dry spring season, develop young daughter cladodes that behave like C3 plants, with daytime stomatal opening and water loss. In contrast, wild Opuntia are less affected because daughter cladodes do not develop on them under extreme drought conditions. The main objective of this work is to evaluate the effects of the number of daughter cladodes on gas exchange parameters of mother cladodes of Opuntia ficus-indica exposed to varying soil water contents. METHODS: Rates of net CO2 uptake, stomatal conductance, intercellular CO2 concentration, chlorophyll content and relative water content were measured in mature mother cladodes with a variable number of daughter cladodes growing in spring under dry and wet conditions. KEY RESULTS: Daily carbon gain by mother cladodes was reduced as the number of daughter cladodes increased to eight, especially during drought. This was accompanied by decreased mother cladode relative water content, suggesting movement of water from mother to daughter cladodes. CO2 assimilation was most affected in phase IV of CAM (late afternoon net CO2 uptake) by the combined effects of daughter cladodes and drought. Rainfall raised the soil water content, decreasing the effects of daughter cladodes on net CO2 uptake by mother cladodes. CONCLUSIONS: Daughter cladodes significantly hasten the effects of drought on mother cladodes by competition for the water supply and thus decrease daily carbon gain by mother cladodes, mainly by inhibiting phase IV of CAM.