Both quinoa production and consumption worldwide have been growing exponentially. As an example, quinoa cultivation has increased from seven countries in 1980 to 123 countries in 2018. Despite the rapid expansion of this crop, countries such as Peru and Bolivia continue to supply 74% of the world market for this grain. Nevertheless during the last decade the production prices of quinoa collapsed, while the yields remained low, so it is necessary to propose local strategies that are based on the diversity of cultivars, their incorporation of food and cooperation to favor its consumption (Alandia et al., 2020).

This situation is gaining importance in Colombia, where recent studies lacked diversity in the genetic material present and cultivated in the country (Guerrero-López, 2018). However, because of non-incorporation of quinoa in the annex list of the Convention on Biological Diversity held in Rio de Janeiro, its seeds began to be used in different parts of the world, which changed its biological behavior when established under different agroecological environments (soil, rainfall, temperature, and altitude; Ruiz et al., 2014). This situation also occurs in Colombia, where there are more than 30 accessions different in genetic, morphological, and productive attributes (Manjarres-Hernández et al., 2021).

Quinoa diversity analyses have been conducted under controlled or cold climate conditions and lack information related to compositional characteristics of the grains (Torres et al., 2000; Delgado et al., 2009; García-Parra et al., 2019). For this reason, Reguera et al. (2018) highlights the importance of evaluating quinoa cultivars through the interaction between environment and genotype for more information, leading to a complete analysis of this species. However, the information that reports the effect of the environment on the compositional characteristics of quinoa grains is clear and mainly on lipid, protein, starch, fiber, and bioactive compound structures (Miranda et al., 2013; Curti et al., 2019; Matías et al., 2022).

One of the climatic factors with greatest impact on the productive performance of plants is altitude, as it brings together climatic elements associated with radiation and temperature (Haeberli and Alean, 1985). Temperature is crucial for active and passive transport between source and dump organs, as well as their structuring in fruits (Smith et al., 2018). Such a situation is vital in photoassimilate transport rate, as well as in transport pathway, for which plant seeds conform into protein, lipid, and starchy structures and depend on individual genetic basis and the environment, as well as the transport pathway developed in relation to the movement of photoassimilates, as highlighted by Lee et al. (2019) in rice plants.

In this sense, Thiam et al. (2021) present results of the environment-genotype interaction in quinoa cultivars and find a strong influence of climatic elements on grain yield parameters. The previous results were similar to those obtained by Granado-Rodríguez et al. (2021), who determined that the environment-genotype interaction modifies the nutritional response of quinoa grains. Also, García-Parra et al. (2021) made the first report on the effect that genetic variation (cultivars) has on the quantity and secondary structure of starch (amylose and amylopectin ratio) and proteins (β-sheets 1 and 2, random coil α-elice, and β-turns 2 and 3) present in quinoa grains.

This phenomenon results from the biological and physicochemical responses of quinoa grains (Bertero et al., 2004; Stikic et al., 2012); however, most of the studies have advanced in the effect that different industrial processes have on the grains, but they forget the influence the environment-cultivar interaction has by contributing in the chemical and phytochemical characteristics of quinoa seeds (Antognoni et al., 2021; Granado-Rodríguez et al., 2021). For this reason, this study aims to find an effect of agroclimatic conditions on the physicochemical and rheological characteristics of different quinoa cultivars in the Department of Boyacá-Colombia.

Quinoa is often compared to cereals, since its characteristics are similar in composition and use. However, its agro-industrial potential is not yet reached under new cultivation regions, with deficiencies in research that relates primary production with the agro-industrial characteristics of grains compared mainly to crops such as maize, rice, and wheat (Stikic et al., 2012). The establishment and adaptation of quinoa cultivars to these new edaphoclimatic contexts can result in changes in nutritional properties of quinoa grains, which are associated with variations in the genotype, the environment, and their interaction (Granado-Rodríguez et al., 2021).

As previously described, the results show a changing composition in the proximal allocation of seeds, which ratifies the effect that climate and its relationship with the genetic variation of the species have on the potential use of grains. This is evidenced in other cultivars and under tropical and extratropical conditions compared by Reguera et al. (2018) who highlight the effect of climate, soil, water, fertilization, establishment, and management practices, as well as quinoa genotype. Thus, the production of four quinoa cultivars at two different sowing times affected the protein concentration in the grains, which is attributed to changes in precipitation and temperature during the test periods (Präger et al., 2018).

In quinoa, the effect of edaphic nitrogen and radiation on the characteristics of quinoa grains has been reported (Bascuñán-Godoy et al., 2018; Eustis et al., 2020), and is mainly attributed to the effect of temperature environmental and the function of mineral elements on the exchange rate between CO₂ and water vapor, its incorporation in the carbon fixation phase and the ability to produce triose phosphate molecules for storage as starch in the chloroplasts or transported to the cytosol as hexoses that initiate the structuring of the reserve organs, mainly the perisperm where the greatest part of carbohydrates (Lemoine et al., 2013). In addition, it has been reported that in the anthesis phase, the transport of hexoses and amino acids to ovule cells is regulated by the characteristics of the abiotic environment, being able to change from symplastic to apoplastic transport, determining the compositional characteristic of the grains and being able to modify the concentration of proteins, carbohydrates, lipids, and water in grains (Zúñiga-Sánchez et al., 2017).

The secondary structure of quinoa has not been well-studied by deconvolution spectroscopy. However, García-Parra et al. (2021) and Roa-Acosta et al. (2020) identified structures such as β-sheet, random coil, α-helix, β-turns, and β-type, between the bands 1,600 and 1,700 cm⁻¹, being able to change the intensity of the peaks of these structures because of the effects of genetic diversity, agro-industrial processing, or climatic factors during primary production, as manifested for other species (Nawrocka, 2013; Li et al., 2020; Pezzotti et al., 2021). Table 2 shows that the 1,547 and 1,634 cm⁻¹ bands associated with amides II and I present a higher peak intensity in the cold and warm climates, responding to the influence generated by the environmental temperature and hydric status on the speed of gas exchange, transport of substances, and their discharge in storage organs; such a phenomenon is very variable in plants that present C3 metabolism (Sonnewald et al., 2020), because the stomatal opening determines the speed of multiple physiological activities and accumulation of proteins in the seeds. A similar situation occurs in the 1,015 cm⁻¹ band associated with starch, the highest proportion of which was found in cultivars in cold climates, which is a natural response of the species, since its ideal abiotic condition brings together climatic elements of cold zones (Torres et al., 2000).

Although some studies report that the amount of proteins is not affected by edaphoclimatic factors such as temperature, edaphic fertilization, and even growth season (Lesjak and Calderini, 2017; Präger et al., 2018; García-Parra et al., 2020), it has been found that the molecular organization of proteins can change, affecting their organization and functionality, as affirmed by Wang et al. (2020), who identified four secondary protein structures in six varieties of Chinese quinoa present in bands like those found in this study. However, both the intensity of the peaks and the band where the characteristic bonds of each structure vibrate or flex may change because of the variety used, characteristics of the production area, and management practices in primary production.

As production climate is a determining factor in the structural configuration of proteins, it is necessary to emphasize that not only the agro-industrial processes developed with the grains can change their techno-functional characteristics but also that performance during the growth and development of the grains can modify their compositional characteristics. In this sense, Wolkers et al. (1998) found that the maturation speed in seeds changes the intensity of the peaks of the secondary protein structure, especially in α-helix and β-sheet, which increased when presenting slower maturation periods, becoming an additional factor of evaluation for the grains of different quinoa cultivars together with the climate where they are grown and the speed of maturation.

However, the changes were not only given in the protein structure but also in the starchy conformation of the grains. Therefore, the band corresponding to 1,047 cm⁻¹ is associated with crystalline structures, the 1,022 cm⁻¹ is attributed to amorphous structures in starch, which is the reason the different ratios between amorphous and crystalline structures influence the order in this fraction of the grains, while the 995 cm⁻¹ region is given to bending and vibration of C-OH bonds (Ahmed et al., 2018). However, the functionality of starch in food matrices has been attributed mainly to species, ignoring the effect of climate as reported by Labuschagne et al. (2007) and Yu et al. (2015) when establishing its effect on amylose/amylopectin ratio in grains. Thus, an example of its importance lies in its use to obtain gelled or translucent films, given that a high crystallinity index is required, while its incorporation into nutritional matrices requires starchy structures with low crystallinity indexes, since it reduces the retrograde phenomenon (Selma-Gracia et al., 2020).

Among the cultivars in diverse climates, those grown in warm climates showed, on average, higher peak viscosity compared to those grown in cold and temperate climates. This phenomenon can be attributed to environmental conditions, even though starch biosynthesis is regulated by genetic expressions of the species. It has been reported, that changes in photosynthetic rate due to effects on the inhibition of electron transport in photosystems during fruit growth periods alter the activity of starch-sitases (Tofiño et al., 2006), that of GBSSI that synthesizes amylose (Raynaud et al., 2016). In addition, inactivation of the enzyme ADP-glucose pyrophosphorylase induces increase in amylopectin content and transport of proteins associated with perisperm plastids, changing the kinetic properties of the kernels (Tetlow et al., 2004), since proteins cannot gelatinize like starch.

In this sense, the same cultivars with highest viscosity peak presented highest values in the setback, which is associated with high amylose contents in the perisperm matrix of quinoa grains, obtaining gels with higher water absorption capacity within the free linear and helical structures, as hydrogen bonds capture water molecules by reorganizing and generating gels with high viscosity peaks (Solarte-Montúfar et al., 2019).

The results obtained in this investigation support the idea associated with responses that cultivars can manifest as a result of their interaction with photo-thermal and precipitation conditions to determine the content of proteins, carbohydrates and lipids, as well as their structural organization, thus obtaining grains with different potential for use in the food industry by proper selection of cultivars and growth conditions throughout the primary production cycle.

In this sense, despite the fact that the secondary structure of proteins and starch are considered fingerprints for differentiation of the composition of quinoa grains, it is important to highlight that the intensity of peaks is modified by the effect of conditions where the cultivars are established, since this study reports the first advances of the significant differences that the secondary structures present through the FT-IR deconvolution in the intensity of the peaks associated with β-sheet, α-helix and β-turns, Random coil.

In structural organization of starch, bands associated with crystalline and amorphous structures of the grains were significantly changing. However, the 1,075 cm⁻¹ band was shown as a region in the spectrum that is not affected by the two factors evaluated and is proposed as an attribute for monitoring and quality of quinoa products. In this sense, the Soracá cultivar presented the highest ratio of peaks 1,047/1,022 cm⁻¹, allowing films with better gelation and translucent characteristics, while the Puno, Nariño, and Titicaca cultivars presented the lowest values for this ratio, making them important for its incorporation in food matrices where the retrogradation phenomenon is avoided because of low crystallinity index.

From the data, it was possible to find that a satisfactory adjustment to the power model is presented, determining the dispersion flow index of the treatments and facilitating the description of their rheological behavior. In addition, it is evident that the treatments presented a dilatant behavior before heating, but that the dispersion of the treatments presented a pseudoplastic behavior after the thermal treatment. In this sense, the behavior of the consistency index after heating/cooling was relevant, since it was affected by the treatments and was mainly associated with the composition of starch granules in the amorphous and crystalline regions, as well as its relationship with the protein for the formation of gels, generating a marked differentiation between the established treatments.

The original contributions presented in the study are included in the article/supplementary material, further inquiries can be directed to the corresponding author/s.

MG-P, DR-A, and JB-G: conceptualization. DR-A, MG-P, JH-C, and HV-C: methodology. MG-P and DR-A: software and investigation. MG-P and JH-C: formal analysis. HV-C: resources. All authors have read and agreed to the published version of the manuscript.

The authors express gratitude to Minciencias' (Ministerio de Ciencia, Tecnología e Innovación, Before Colciencias) invitation for bid Nr. 779/2017. We are also grateful to the Boyacá Department Government and Universidad del Cauca.

The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.

All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.