Secondary Metabolism in Amaranthus spp. — A Genomic Approach to Understand Its Diversity and Responsiveness to Stress in Marginally Studied Crops with High Agronomic Potential

Cultivated amaranths are crops with an unrealized agronomical potential despite their high nutritional value and nutraceutic properties of their seeds and/ or leaves. They tolerate growing conditions unsuitable for cereals, and are tolerant to biotic aggressors. Several Amaranthus species are abundant of sources of secondary metabolites, mostly phenylpropanoids, predominantly in seeds and leaves, many of which may confer health benefits associated with their antioxidant properties. They could also act as defensive compounds against predators or pathogens. Recent biochemical and molecular approaches partly defined the mechanisms responsible for grain amaranth´s tolerance against biotic stress. However, the role played by secondary metabolites in (a)biotic stress amelioration in amaranth is practically unknown. Our group has identified several genes coding for enzymes involved in secondary metabolism pathways in A. hypochondriacus, in addition to related regulatory transcription factors. More than 50% of these genes involve the phenylpropanoid pathway. In this chapter, the role played by this pathway in (a)biotic stress amelioration in plants will be briefly reviewed, followed by an examination of its involvement in the conferral of nutraceutic properties to amaranth plants. A description of the progress obtained so far regarding the characterization of phenylpropanoid genes in grain amaranth will close this chapter

Metabolic and enzymatic changes associated with carbon mobilization, utilization and replenishment triggered in grain amaranth (Amaranthus cruentus) in response to partial defoliation by mechanical injury or insect herbivory

Background: Amaranthus cruentus and A. hypochondriacus are crop plants grown for grain production in subtropical countries. Recently, the generation of large-scale transcriptomic data opened the possibility to study representative genes of primary metabolism to gain a better understanding of the biochemical mechanisms underlying tolerance to defoliation in these species. A multi-level approach was followed involving gene expression analysis, enzyme activity and metabolite measurements. Results: Defoliation by insect herbivory (HD) or mechanical damage (MD) led to a rapid and transient reduction of non-structural carbohydrates (NSC) in all tissues examined. This correlated with a short-term induction of foliar sucrolytic activity, differential gene expression of a vacuolar invertase and its inhibitor, and induction of a sucrose transporter gene. Leaf starch in defoliated plants correlated negatively with amylolytic activity and expression of a β-amylase-1 gene and positively with a soluble starch synthase gene. Fatty-acid accumulation in roots coincided with a high expression of a phosphoenolpyruvate/phosphate transporter gene. In all tissues there was a long-term replenishment of most metabolite pools, which allowed damaged plants to maintain unaltered growth and grain yield. Promoter analysis of ADP-glucose pyrophosphorylase and vacuolar invertase genes indicated the presence of cis-regulatory elements that supported their responsiveness to defoliation. HD and MD had differential effects on transcripts, enzyme activities and metabolites. However, the correlation between transcript abundance and enzymatic activities was very limited. A better correlation was found between enzymes, metabolite levels and growth and reproductive parameters. Conclusions: It is concluded that a rapid reduction of NSC reserves in leaves, stems and roots followed by their long-term recovery underlies tolerance to defoliation in grain amaranth. This requires the coordinate action of genes/enzymes that are differentially affected by the way leaf damage is performed. Defoliation tolerance in grain is a complex process that can’t be fully explained at the transcriptomic level only

Importancia agroecológica del coyul (Acrocomia mexicana Karw. ex Mart.)

Acrocomia mexicana Karw. ex Mart. es una especie que se extiende desde el Pacífico mexicano hasta Centro y Sudamérica. El fruto (coyul) es apreciado desde la época prehispánica, sin embargo, su consumo ha disminuido frente a otros frutos nativos y así como de los introducidos con mayor aceptación en el mercado. Se realizó un estudio en San Blas, Nayarit, México, en el que se llevó a cabo la caracterización de un agroecosistema típico donde se explotan poblaciones silvestres y cultivadas de coyul de forma tradicional y se resaltó la importancia económica, ecológica y cultural de la especie. Asimismo, se realizó la caracterización fisicoquímica del fruto; la pulpa y la semilla son fuente principalmente de grasas. De esta especie se pueden obtener subproductos tales como aceites comestibles, alimento para ganado, artesanías, entre otros; igualmente se exponen alternativas de uso llevadas a cabo por otras regiones y países que puedan conducir a conservar espacios ecológicos y culturales

The novel and taxonomically restricted Ah24 gene from grain amaranth (Amaranthus hypochondriacus) has a dual role in development and defense

Grain amaranths tolerate stress and produce highly nutritious seeds. We have identified several (a)biotic stress-responsive genes of unknown function in Amaranthus hypochondriacus, including the so-called Ah24 gene. Ah24 was expressed in young or developing tissues; it was also strongly induced by mechanical damage, insect herbivory and methyl jasmonate and in meristems and newly emerging leaves of severely defoliated plants. Interestingly, an in silico analysis of its 1304 bp promoter region showed a predominance of regulatory boxes involved in development, but not in defense. The Ah24 cDNA encodes a predicted cytosolic protein of 164 amino acids, the localization of which was confirmed by confocal microscopy. Additional in silico analysis identified several other Ah24 homologs, present almost exclusively in plants belonging to the Caryophyllales. The possible function of this gene in planta was examined in transgenic Ah24 overexpressing Arabidopsis thaliana and Nicotiana tabacum plants. Transformed Arabidopsis showed enhanced vegetative growth and increased leaf number with no penalty in one fitness component, such as seed yield, in experimental conditions. Transgenic tobacco plants, which grew and reproduced normally, had increased insect herbivory resistance. Modified vegetative growth in transgenic Arabidopsis coincided with significant changes in the expression of genes controlling phytohormone synthesis or signaling, whereas increased resistance to insect herbivory in transgenic tobacco coincided with higher jasmonic acid and proteinase inhibitor activity levels, plus the accumulation of nicotine and several other putative defense-related metabolites. It is proposed that the primary role of the Ah24 gene in A. hypochondriacus is to contribute to a rapid recovery post-wounding or defoliation, although its participation in defense against insect herbivory is also plausible

Grain amaranths are defoliation tolerant crop species capable of utilizing stem and root carbohydrate reserves to sustain vegetative and reproductive growth after leaf loss.

Tolerance to defoliation can be defined as the degree to which productivity is affected by photosynthetic area reduction. This trait was studied in grain amaranth (Amaranthus cruentus and A. hypochondriacus), which are considered to be a highly defoliation-tolerant species. The physiological and biochemical responses to increasing levels of mechanical leaf removal up to total defoliation were quantified. Tolerance appeared to be dependent on various factors: ( i) amount of lost tissue; (ii) mechanics of leaf tissue removal; (iii) environment, and (iv) species tested. Thus, grain amaranth was found to be a highly tolerant species under green-house conditions when leaf tissue loss was performed by gradual perforation. However, tolerance was compromised under similar conditions when defoliation was done by gradual cutting of the leaf. Also tolerance in completely defoliated plants tended to decrease under field conditions, where differences between A. cruentus and A. hypochondriacus were observed. All non-structural carbohydrate (NSC) levels were reduced in stems and roots of totally defoliated amaranths one day after treatment. Such depletion probably provided the carbon (C) resources needed to sustain the early recovery process in the absence of photosynthetic capacity. This was corroborated by shading of intact plants, which produced the same rapid and drastic reduction of NSC levels in these tissues. These results emphasize the role of stored NSCs, particularly starch, in buffering the impact of severe defoliation in amaranth. The fall in sucrose synthase and cell wall invertase activity observed in stems and roots soon after defoliation was consistent with their predicted shift from sink to source tissues. It is concluded that mobilization of C stores in stems and roots, is a physiologically important trait underlying tolerance to defoliation in grain amaranth