Abstract
Resistance to
Introduction
Maize,
Comparative studies between maize and teosintes have shown that the resistance mechanisms for herbivorous insects are stronger on teosinte than on maize, because of changes in life history, domestication process, and agronomic selection (Rosenthal and Welter 1995; Rosenthal and Dirzo 1997; Takahashi et al. 2012; Szczepaniec et al. 2013; Bernal et al. 2015; Moya-Raygoza 2016). Regarding this, perennial teosintes (
Nutritional quality of the host plant is one of the main factors involved in the development and fecundity of insects: the content of carbon, nitrogen, and defensive compounds (alkaloids, non-protein amino acids, glycosides, cyanogenic glucosinolates, terpenoids, and phenols) are important (Awmack & Leather 2002). In the assessment conducted by Takahashi et al. (2012),
In Mexico, there are approximately 65 maize races (Serratos-Hernández 2012; Santillán-Fernández et al. 2021) and six species and subspecies of teosintes (Sánchez-González et al. 2018). Tuxpeño (De La Rosa-Cancino et al. 2016), Elotillo, Nal Tel (dos Santos et al. 2018), Uruapeño (Farias-Rivera et al. 2003), and Zapalote Chico (Widstrom et al. 2003; Nuessly et al. 2007; de Oliveira et al. 2018; Michereff et al. 2018; Crubelati-Mulati et al. 2019) have been documented as maize races with resistance or tolerance to
Materials and methods
Insect collection and rearing
To maintain
Host plants
Seeds of Zapalote Chico maize and
Development parameters
Feeding groups were integrated with three
Fecundity parameters
When the adults emerged, the sex of each individual was determined to obtain the proportion of females and males (total females or males/total individuals) for each feeding group, and they were immediately confined in Kraft paper bags to promote mating. Once the oviposition process began, the moths were changed from the bags daily to remove the egg masses and to count the number of eggs produced. The eggs produced per day were counted with a stereoscope microscope Carl Zeis™ Stemi™ DV4 (ZEISS Group, Göttingen, Germany). The duration of oviposition, the number of eggs per egg mass, the number of eggs per female, and the total eggs per host plant type were determined.
Data analysis
All data were analyzed using the independent samples t-test and the Mann-Whitney U test as statistical analysis, with a confidence level of 95%. Differences were analyzed by host plant type and were considered significant when P ≤ 0.05. Student's t-test was used for variables with normal distribution and the Mann-Whitney U-test for variables with non-normal distribution, as indicated by McKnight and Najab (2010). The normality of data was determined by Kolmogorov-Smirnov and Shapiro-Wilk tests, and the homogeneity of variances by Levene's test. For data presented as a percentage, before the statistical analysis, an arcsine transformation [√(x/100)] was used. In addition, to determine the relationship between development parameters of the different stages of
Results
Development parameters
The larval survival of
Feeding of
Development parameters (mean ± sd) of males and females of S. frugiperda fed on native maize and teosinte from Oaxaca, Mexico.
Values with the same letter in a row do not differ significantly (P>0.05): 1Mann-Whitney U test (U = 3,390.0, p = 0.58); 2Mann-Whitney U test (U = 3,451.0, n = 35, p = 0.72); *3Mann-Whitney U test (U = 3,314.00, n = 35, p = 0.43); *4t-test (t = -0.84, df = 22, p = 0.41); $5t-test (t = -0.97, df = 17, p = 0.33); $t-test(t = 0.38, df = 167, p = 0.70); $t-test (t = 3.17, df = 167, p = 0.00); $8t-test (t = 0.57, df = 22, p = 0.57); 9Mann-Whitney U test (U = 56.00, n = 35, p = 0.36); †1Mann-Whitney U test (U = 2,507.00, n = 35, p = 0.06); †βMann-Whitney U test (U = 2,928.50, n = 35, p = 0.69); †³Mann-Whitney U test (U = 2,629.00, n = 35, p = 0.15); †4t-test (t = -0.84, df = 22, p = 0.41); †5t-test (t = -0.07, df = 154, p = 0.94); †6t-test (t = 0.59, df = 154, p = 0.55); †7t-test (t = 1.92, df = 154, p = 0.06); †8t-test (t = -0.57, df = 22, p = 0.57); †Mann-Whitney U test (U= 56.00, n = 35, p = 0.36); n: number of larvae that constitute the sample size.
Fecundity parameters
There were no statistically significant differences (P > 0.05) between the fecundity parameters of
Correlations among parameters of development and fecundity
Regarding males, positive and significant correlations were determined between the durations of all developmental stages. The prolonged duration of the larval stage led to an extended pupal stage, so the duration from hatching to adult emergence was also prolonged (Table 3). Adult emergence was positively correlated with larval survival and pupal formation. Adult lifespan was negatively correlated with larval survival and adult emergence, and positively correlated with larval weight. Larval weight was negatively correlated with larval survival, and maximum larval weight was negatively correlated with larval survival and adult emergence, but positively correlated with larval weight. Possibly, the heavier larvae had problems completing pupal formation. The Fitness Index was positively correlated with larval survival and pupal weight, but negatively correlated with the duration of developmental stages. Negative correlations between the Fitness Index and the duration of larval, pupal, and from hatching to adult emergence stages, are congruent with the formula, because the longer duration of developmental stages, the lower the value of the Fitness Index. An increase in pupal weight led to an increase in the performance of male offspring.
Concerning females, a similar pattern for the duration of developmental stages, adult emergence, and adult lifespan was determined (Table 4). In addition, the prolonged duration of developmental stages was negatively correlated with larval weight, maximum larval weight, and pupal weight. Maximum larval weight was positively correlated with larval and pupal weight. The Fitness Index was positively correlated with larval survival, adult emergence, and pupal weight, but negatively correlated with the duration of developmental stages. Regarding the fecundity parameters of
Fecundity parameters (mean ± SEM) of S. frugiperda females fed on native maize and teosinte from Oaxaca, Mexico.
Values with the same letter in a row do not differ significantly (P > 0.05): †1t-test (t = -0.68, df = 18), †βMann-Whitney U test (U= 165,990.50, n = 35, p>0.05), †³t-test (t=0.67, df = 18), †4t-test (t=0.287, df=18).
Pearson correlation coefficient among development parameters of S. frugiperda males.
The correlation is significant at the 0.05 level
The correlation is significant at the 0.01 level. LS: larval survival, PF: pupal formation; AE: adult emergence; DLS: duration of the larval stage; DPS: duration of the pupal stage; HAE: hatching to adult emergence; DAS: duration of the adult stage; LW: larval weight; MLW: maximum larval weight; PW: pupal weight; rL: Fitness Index.
Pearson correlation coefficient among development parameters and fecundity of S. frugiperda females.
The correlation is significant at the 0.05 level
The correlation is significant at the 0.01 level. LS: larval survival, PF: pupal formation; AE: adult emergence; DLS: duration of the larval stage; DPS: duration of the pupal stage; HAE: hatching to adult emergence; DAL: duration of the adult stage; LW: larval weight; MLW: maximum larval weight; PW: pupal weight; rL: Fitness Index; DO: duration of oviposition; E/EM: eggs per egg mass; E/F: eggs per female; E/H: eggs per host plant type.
Discussion
In this study, it was hypothesized that there would be an increase or decrease in the development parameters and fecundity of
The developmental pattern of
No statistically significant differences were determined between the Fitness Index for both sexes and the host plant types but it was higher on teosinte than on native maize, and the highest value was determined for males. These results were consistent with the pattern observed in the pupal weight because the highest value was determined for males, and the difference was greater on teosinte. Similar data were reported by Nogueira et al. (2019) in their evaluation with Brazilian maize races that were tolerant to
Regarding the fecundity of
In Pearson correlation analyses, for both males and females, the Fitness Index was negatively correlated with the duration of developmental stages but positively correlated with the pupal weight. Similar results were reported by Boregas et al. (2013) on
Conclusions
Our results demonstrated that larval feeding on native maize and teosinte did not affect the development parameters and fecundity of
Origin and funding
The present work derived from the research line of Protección y Producción Vegetal developed at the IPN-CIIDIR Unidad Oaxaca. The financial support was provided by CONACYT-Problemas Nacionales N° 1119 project and IPN-SIP (Secretaría de Investigación y Posgrado) projects: 20200821 and 20211461.
Author contribution
Erika Padilla-Cortes: performed the collection, rearing and handling of insects, carried out the experiment and collected the data, realized the analysis and interpretation of results, and wrote the draft and the final manuscript.
Laura Martínez-Martínez: helped in the collection, rearing and handling of insects, supervised the experiment, and provided suggestions to improve the manuscript.
José Luis Chávez-Servia: provided the maize and teosinte seeds, contributed to the analysis and interpretation of results, and provided suggestions to improve the manuscript.
Conflict of Interest
The authors declare that they have no conflict of interest.
Footnotes
Acknowledgements
We thank Roselia Jarquín López for her help in the insect field collection, Prisciliano Diego Flores for collecting the maize seeds, José Antonio Sánchez García and Celerino Robles Pérez for the facilities to use their equipment. We also thank Robert Leavitt and Timothy O'Hara for reviewing the English translation of the manuscript. The first author thanks to Consejo Nacional de Ciencia y Tecnología (CONACYT) for the scholarship provided to carry out her Master's studies. We appreciate the funding of the CONACYT-Problemas Nacionales N° 1119 project and IPN – SIP (Secretaría de Investigación y Posgrado) projects: 20200821 and 20211461.
