1. Introduction
Chitosan is a natural polysaccharide derived from the deacetylation of chitin and is mainly obtained from the exoskeletons of crustaceans and insects [
1,
2]. The process of alkaline deacetylation is inexpensive compared to the enzymatic process due to its low-cost for industrial production; it uses NaOH at 40–50% to eliminate more than 80% of the acetyl groups to obtain
N-acetyl-
d-glucosamine in β-1,4-
d-glucosamine (CS) [
3]. In the search for alternatives to enable a move towards more sustainable agriculture, CS is a good option because it is biodegradable, biocompatible, and non-toxic to humans [
4,
5]. The wide availability of CS has meant that it has been tested for use in agriculture since the 1980s [
3]. Currently, CS is being studied with regard to the administration of pesticides, fertilizers, and growth regulators in order to increase the efficiency and reduce the number of applications of these conventional products [
4,
6]. Due to its properties, CS promotes plant growth and protects plants from biotic and abiotic stress [
7,
8,
9]. The chitosan signaling mechanism includes specific receptors and secondary messengers, such as reactive oxygen species (ROS), hydrogen peroxide, calcium, nitric oxide (NO), and phytohormones that induce physiological responses to mitigate biotic and abiotic stress, and promote plant growth [
8]. The positive effects of CS on plants include improvements in physiological mechanisms and growth, as well as an increase in the shelf life of fruits and vegetables [
9,
10]. These effects have been demonstrated in model plants, including tomato, maize, wheat, cucumber, strawberry, and chili peppers, among others [
5,
9,
10].
Poly(acrylic acid) is a synthetic polymer that is used as a chelating, dispersal, flocculating, and adhesive agent in agricultural soils [
11]. It increases soil water availability to aid plant growth [
12] and can be used in the remediation of heavy-metal contaminated soils [
13]. In addition, it is used to form complexes with chitosan and has previously been applied to seeds, substrates, leaves, and on post-harvest fruits, showing a strong capacity for biostimulation [
14]. In tomato plants, the application of chitosan-poly(acrylic acid) complexes increased yield [
15]. In onion plants, chitosan-poly(acrylic acid) hydrogel nanoparticles improved growth rates and yield [
16]. In lettuce plants, chitosan–poly(acrylic acid) complexes were shown to increase the biofortification with selenium [
17]. Recently, biostimulants in crops have been used more frequently in order to improve productivity [
18].
In Mexico, the production of habanero peppers has increased exponentially over the past two decades, increasing from 38.8 tons in 1999 to 20,829.6 tons in 2019 [
19]. It is considered to be an economically important crop due to the current high demand for the consumption of fresh fruits, its use as an ingredient in sauces, as a natural colorant, and its medicinal uses. Habanero peppers are traditionally produced in soil, so information regarding nutrient solutions when grown in a greenhouse is scarce [
20]. Few studies have evaluated different nutritional regimens used in hydroponic systems to produce habanero peppers in Mexico [
20,
21,
22]. Therefore, it is important to carry out studies in Mexico, where findings will help to increase the productivity of the growth of habanero peppers; most of the published studies have focused on capsaicinoids [
23]. Therefore, the objective of this study was to evaluate the effect of foliar applications of chitosan–poly(acrylic acid) complexes (CS–PAA) and two nutrient solutions on the growth and yield of two habanero pepper cultivars under greenhouse conditions.
4. Discussion
Chitosan is used in horticultural crops to stimulate growth and productivity [
9]. However, these responses depend on the degree of deacetylation, molecular weight, and the concentration of chitosan, as well as the plant species and the phenological stage of the plant [
28]. This polymer is synthesized in combination with poly(acrylic acid) to form complexes that have been shown to have a biostimulant effects on plants [
14]. In the present study, we demonstrated that the foliar applications of CS–PAA complexes increased growth and dry biomass parameters (except dry root biomass) of habanero peppers compared to the control. Meanwhile, the effect of the interactions showed that the applications of the CS–PAA complexes improved the height of the Chichen Itza hybrid and increased the total dry biomass of both cultivars. We did not find any published studies that discussed the effect of chitosan, poly(acrylic acid), or CS–PAA complexes on the growth of
C. chinense. Similar results show that chitosan–poly(acrylic acid) hydrogel nanoparticles improved the growth of onion plants [
16]. Of the few studies regarding
Capsicum, Chookhongkha et al. [
29] reported that the application of chitosan to the soil increased the plant height of
Capsicum annuum cultivated in a greenhouse. Meanwhile, Esyanti et al. [
30] showed that the foliar application of chitosan increased the plant height of
C. annuum cultivated in a shade house. In
C. frutescens cultivated in a greenhouse, the foliar application of oligochitosan increased the fresh and dry weight of the buds [
31]. In addition, the foliar application of chitosan was shown to increase the growth of other species such as tomato, cucumber, strawberry, potato, maize, and wheat, as reported by Mukhtar Ahmed et al. [
5] in their review.
Our results showed that the foliar applications of CS–PAA complexes increased fruit number, yield, and TSS value of ripe habanero pepper fruits compared to the control. Meanwhile, the effect of the interactions showed that the foliar applications of CS–PAA complexes increased the number of fruits of both cultivars. Furthermore, foliar applications of CS–PAA complexes improved the TSS value of the green fruits of the Jaguar variety when it was irrigated with the NSA. Some studies have reported that the application of chitosan–poly(acrylic acid) complexes increased the yield of tomato and onion [
15,
16]. Other authors, such as Chookhongkha et al. [
29] reported that the application of chitosan to soil increased the number of fruits per plant and fresh weight of fruits of
C. annuum cultivated in a greenhouse. Meanwhile, Mahmood et al. [
32] reported that the foliar application of chitosan increased the yield and average weight of fruits of
C. annuum cultivated in open fields. Regarding
C. frutescens cultivated in a greenhouse, the foliar application of oligochitosan was shown to increase the fresh weight of fruits [
31]. Meanwhile, He et al. [
25] demonstrated that the pre-harvest application of chitosan oligosaccharides increased the total soluble solids values of strawberry fruits as shown in the results reported here.
The biostimulant effect of CS–PAA complexes is probably due to the involvement of chitosan in the regulation of nitrogen and carbon metabolism [
33,
34]. This is supported by previous studies that report that chitosan increases the activity of enzymes such as fructose-1,6-bisphosphatase, sucrose phosphate synthase, sucrose synthase, phosphoenolpyruvate carboxylase, pyruvate dehydrogenase, malate dehydrogenase, nitrate reductase, glutamate synthase, glutamine synthetase, among others [
5,
33,
34,
35,
36]. Similarly, El-Tanahy et al. [
37] and Dwivany Fenny et al. [
38] propose that the biostimulating effect of chitosan on plant growth is due to the primary aliphatic amino groups of the polymer, which provide a source of nitrogen. This corresponds with the findings of Ravi Kumar [
39], who reported that the structure of this polymer can contain up to 6.89% nitrogen. Another hypothesis is that chitosan increases the net photosynthetic rate, which leads to better plant growth and development [
5,
36]. Whether chitosan-specific receptors exist or not is unknown as they have not been clearly elucidated [
33]. So far it is known that the chitosan signaling pathways in plants are calcium, NO, ROS, phytohormones and hydrogen peroxide [
8].
Regarding nutrient solutions evaluated, the NSA increased the stem diameter and the yield of habanero peppers compared to the NSB. Meanwhile, the effect of the interactions showed that the NSA increased the yield and the TSS value of the green fruits of the Chichen Itza hybrid. These responses are due to the relationship between potassium and calcium in the nutrient solution. Potassium and calcium are two cations that present antagonism [
40], so an unbalanced ratio of these cations in the nutrient solution can affect crop yield. Hernández-Pérez et al. [
26] reported that the relationship between potassium and calcium in the nutrient solution is associated with higher yield and sugar concentration in the tomato fruits with the optimal balance being between 0.82 and 0.85, while if this value is above 1.0, yield the sugar concentration then decreases drastically. This coincides with our results as the NSA and NSB present a relationship between potassium and calcium with values of 0.83 and 1.2, respectively. Among the cultivars evaluated, the Chichen Itza hybrid obtained a higher fruit yield than the Jaguar variety. This shows the genetic potential per se of the hybrid compared to the improved variety. The use of hybrid seeds is a good alternative to improve crop yields due to the exploitation of heterosis [
41].
5. Conclusions
Our results showed that the foliar applications of CS–PAA complexes improved plant height, stem diameter, total dry biomass, number of fruits, and yield of habanero peppers compared to the control. Meanwhile, the effect of the interactions showed that foliar applications of CS–PAA complexes increased the total dry biomass and the number of fruits per plant of both cultivars. Regarding nutrient solutions, nutrient solution A increased the yield of the Chichen Itza hybrid compared to the nutrient solution B, this was due to it presenting a better balance of potassium and calcium. Meanwhile, the best cultivar was the Chichen Itza hybrid, because it obtained a higher yield than the Jaguar variety, this was due to the exploitation of heterosis. This research offers advances in the use of CS–PAA as a biostimulant to improve the genetic potential of habanero pepper cultivars. It also provides information on the management of nutrient solutions in the crop of habanero peppers in greenhouses, which is very scarce. This could give guidelines for more researchers to address this research topic.