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Agrociencia

versão On-line ISSN 2521-9766versão impressa ISSN 1405-3195

Agrociencia vol.52 no.3 Texcoco Abr./Mai. 2018

 

Crop sciencie

Phisiological and agronomic traits of hard wheat CIRNO variety C2008 confirm its genetic stability

Leandris Argentel Martínez1  2 

Jaime Garatuza Payán1  * 

María Magdalena Armendáriz Ontiveros1 

Enrico Yépez González1 

Juan Manuel Garibaldi Chávez2 

José Eliseo Ortiz Enriquez3 

Jorge González Aguilera4 

1Instituto Tecnológico de Sonora. 5 de febrero, 818 Sur, Ciudad Obregón, Sonora, México. CP: 85000.

2Universidad de Granma, Carretera a Manzanillo, km 17 ½ Peralejo, Bayamo, Cuba. CP: 85100.

3Campo Experimental “Norman Borlaug”. Carretera Norman E. Bouleaug-INIFAP. Ciudad Obregón, Sonora, México.

4Embrapa Trigo, Rodovia BR 285, Km 294, Passo Fundo - RS. Brasil. Caixa Postal 451 99001-970.


Abstract

Due to the importance of the correct selection of progenitors for an effective genetic improvement in highly productive wheat regions such as the state of Sonora, which contributes 50% of the wheat in Mexico, we conducted a study of morphoagronomic variables of the crystalline wheat CIRNO C2008 variety. The objective was to determine the genetic stability index of the grain yield components in the Valle del Yaqui and Valle del Mayo, during the sowing seasons of 2007-2015. In addition, physiological and agronomic traits were evaluated in the 2015-16 cycle in an area established in CETT-910, as a representative site of the Valle del Yaqui, which were compared with the varietal descriptor and correlated with grain yield to contribute to its physioagronomic characterization. The experimental design was completely randomized and data were taken from different sites of the valleys mentioned in the state of Sonora. The highest grain yield was obtained in the Valle del Yaqui. The variables of plant height and spike length were the components of the yield with the highest index of genetic stability. The variety phenophases occurred properly without affecting the biological cycle. The grain yield in the CETT-910, in the 2015-2016 cycle, was higher than that obtained by the varietal descriptor as a result of good nutritional status and high photosynthesis. The results show that CIRNO C2008 maintains genetic stability of the yield components after eight years of being released for agricultural production in the Valle del Yaqui and Valle del Mayo.

Key words: wheat; CIRNO C2008; physiological characteristics; genetic stability

Resumen

Debido a la importancia de la correcta selección de progenitores para una mejora genética efectiva en regiones altamente productoras de trigo como el estado de Sonora, que aporta 50 % del trigo en México, se realizó un estudio de variables morfoagronómicas de la variedad de trigo cristalino CIRNO C2008. El objetivo fue determinar el índice de estabilidad genética de los componentes del rendimiento de grano en los Valles del Yaqui y del Mayo, en las épocas de siembra de 2007-2015. Además se evaluaron caracteres fisiológicos y agronómicos en el ciclo 2015-16 en un área establecida en el CETT-910, como un sitio representativo del Valle del Yaqui, los cuales se compararon con el descriptor varietal y se correlacionaron con el rendimiento de grano para contribuir a su caracterización fisioagronómica. El diseño experimental fue completamente aleatorizado y se tomaron datos de diferentes sitios de dichos Valles del estado de Sonora. El mayor rendimiento en grano se obtuvo en el Valle del Yaqui. Las variables altura de la planta y la longitud de la espiga fueron los componentes del rendimiento con mayor índice de estabilidad genética. Las fenofases de la variedad transcurrieron debidamente sin afectar el ciclo biológico. El rendimiento de grano, en el CETT-910 en el ciclo 2015-2016, fue superior al obtenido por el descriptor varietal como resultado de buen estado nutricional y alta fotosíntesis. Los resultados muestran que CIRNO C2008 mantiene estabilidad genética de los componentes del rendimiento tras ocho años de liberada para la producción agrícola en los Valles del Yaqui y del Mayo.

Palabras clave: trigo; CIRNO C2008; caracteres fisiológicos; estabilidad genética

Introduction

Multiple plant species, including cereals, and wheat in particular, undergo a change in their morphological and physiological traits that correlate with the genetic stability of agricultural yield (Monaco et al., 2014). This situation may be due to the adverse effect of biotic or abiotic factors, or both, modifying the genotype-environment interaction and making vulnerable the genetic-productive potential of the varieties vulnerable (Silva et al., 2016). Such vulnerability imposes the need to perform the monitoring of the germplasm available based on physiological and agronomic characteristics, when there are varieties established in different regions and for a long time (Lopes et al., 2015).

The use of performance indicators to examine the genetic stability of commercial varieties with years and sites of exposure to the environment allows to identify characters for improvement programs when they still show adaptation to edaphoclimatic conditions, such as resistance or tolerance to biotic and abiotic stresses and that still express their productive genetic potential (Solís et al., 2016).

Due to the importance of wheat cultivation for food, and the Valles del Yaqui and del Mayo, in the Sonora state, Mexico, contributing 50 % of the national production of wheat (SAGARPA, 2015), the objective of the present study was to evaluate the morphological, physiological and agronomic characters of the hard wheat variety CIRNO C2008 to determine the genetic stability index of grain yield and contribute to its physiological and agronomic characterization, after eight years of being released for extensive agricultural use.

Materials and Methods

Location of the experimental area

The research was carried out in the Valles del Yaqui and del Mayo, as representative sites of highly productive regions of irrigated wheat in Mexico, where about 160 and 97 thousand ha, respectively, are sown in the Valle del Yaqui (27° 30' N; 110° 20' W); and Valle del Mayo (26º 41' N; 109º 30' W), state of Sonora.

The main crop in these valleys is wheat with a surface irrigation system whose water comes from the Álvaro Obregón and Adolfo Ruiz Cortinez dams (SAGARPA, 2015). The air temperature in these valleys has minimum and maximum levels of 0 and 45 ° C in winter and summer respectively. The predominant soils are classified as compacted clayly and alluvial (Ortiz and del Carmen, 2015), based on the methodology proposed by Soil Taxonomy (Bhattacharyya et al., 2015).

Genetic stability index

We used the methodology proposed by Annicchiarico (1992) to determine the genetic stability index of the grain yield components: plant height, spike length, grain mass and grain yield. The statistical data of grain yield components of the CIRNO C2008 variety were collected from 2007 to 2015 in the Valles del Yaqui and del Mayo where wheat is cultivated, especially the CIRNO C2008 variety (SAGARPA, 2015).

Main characteristics of the CIRNO C2008 variety

The CIRNO C2008 variety is classified as crystalline or hard wheat (Triticum durum L.) and originated from the selection in segregating populations of the SOOTY-9 / RASCON-37 / CAMAYO cross, carried out in the Centro Internacional de Mejoramiento de Maíz y Trigo, CIMMYT (International Center for the Improvement of Maize and Wheat). This variety is directly related to the Átil C2000 variety, which originated from the SOOTY-9/RASCON-37 crossing; its release for cultivation took place in 2008 and is used mainly in the state of Sonora.

This variety grows in the spring, ideal to be cultivated during the autumn-winter cycle under irrigation conditions. Heading occurs from 74-89 d, the physiological maturity at 122 d on average and the length of stems is of 78 cm on average and is classified as a short-size variety; with erect stems and very low or null frequency of plants with flag recurved leaf. Before maturing, the sheath of the flag leaf and the peduncle of the spike present high levels of wax. Its grain yield when released for agricultural extensive use in Sonora was 5600 and 6300 kg ha-1 with two and three auxiliary irrigations, respectively (Figueroa et al., 2010).

Physiological and agronomic study in the 2015-2016 cycle

The CIRNO C2008 variety was planted in 2015-2016 in order to monitor physiological and agronomic characters in an experimental area of 0.48 ha (88.88 x 54.0 m), under open field conditions, in Block 910 of the Instituto Tecnologico de Sonora´s Centro Experimental de Transferencia de Tecnología (Technology Transfer Experimental Center) (CETT), located at 27 ° 22'0.4 '' N and 109 ° 54' 50.6 '' W (UTM coordinates: 607393.24 m E; 3027508.34 m N).

Sowing, fertilization and maintenance of the crop in the 2015-2016 cycle

Sowing was carried out with a planter (ST 16) on December 8, 2015, in three rows on the quarry with a planting density of 138 kg ha-1. Fertilization was based on 300 kg ha-1 of urea + 100 kg ha-1 of monoammonium phosphate (MAP) 11-52-00. In the first auxiliary irrigation, 150 kg ha-1 of urea were applied and the other part was divided in the second and third irrigations at a rate of 75 kg ha-1 each.

The irrigations were at pre-sowing, tillering, heading and grain filling with a mean water depth of 16 cm, and a gross partial standard of Nb = 160 m3 ha-1 in each irrigation until reaching field capacity. The average watering interval was 25 d. For these tasks we followed the technical instruction for this variety (Figueroa et al., 2010).

Control of pests and diseases

The foliage aphid (Schizaphis graminum) was controlled with the pesticide Muralla Max (ia Imidacloprid + Betaciflutrin) (0.20 L ha-1 in the periphery of the plot up to 2.0-3.0 m inwards (edge surface), at the beginning of heading, when the pest was present. The broadleaf weeds were controlled with manual weeding before applying auxiliary irrigations.

Agroclimatic variables during the 2015-2016 crop cycle

During the biological cycle of the crop, the average monthly temperature remained between 17 and 24 ° C (average 18.6 ° C). Monthly precipitation was less than 0.2 mm and relative humidity varied from 50 to 78 % (Figure 1).

Figure 1 Climatic variables (temperature, precipitation and relative humidity) during the cultivation cycle December 2015 - May 2016 at the Technological Transfer Experimental Center of the Technological Institute of Sonora (CETT). 

Variables evaluated in the CETT experiment in the 2015-16 cycle

Phenology of the variety

To study the phenological response we used the decimal scale of comparison by Zadoks (1974), considering the phenophase when more than 50% of the population showed the related characteristics. The time periods of the phenophases were compared with that obtained by the varietal descriptor. The variables studied were the days at the start of tillering, days at the appearance of the first node, days to heading, days of grain filling, and days at physiological maturity.

Normalized difference vegetation index (NDVI)

The normalized difference vegetation index was measured with a portable sensor (Green Seeker, Trimble ™ brand) (Govaerts and Verhulst, 2010), from 15 d after germination in each phenophase until physiological maturity of the grain. In each phenophase we took 30 measurements at 0.60 m high from the crop canopy, according to the sensor reference.

This variable was evaluated to compare in each phenophase the value of the normalized difference vegetation index NDVI (its acronym in English); -1<NDVI>1, whose interpretation can contribute to the rapid and targeted diagnosis of the nutritional conditions of the crop (especially nitrogen) and the possible incidence of stress. Greater values of the NDVI represent a better nutritional status (Inman et al., 2005).

Maximum photosynthesis

The photosynthetic activity was measured in leaves and spikes in the phenophases of heading, flowering and grain filling at biweekly intervals, and was evaluated with a portable system (LI-6400XT, LI-COR, Inc.) that measures CO2 concentration and water vapor through a spectrometer that operates in the infrared spectrum of the electromagnetic radiation (IRGA, gas analyzer in the infrared spectrum, its acronym in English).

Measurements were taken between 1000 and 1100 on sunny days. For this measurement, the three leaves most exposed to direct solar radiation (repetitions per plant) were inserted by their central part into the natural gas exchange light of 3.0 x 2.0 cm. The same was done with the spikes (using the specific camera for such organ), with measurements taken in the central part.

All measurements were made with a luminous intensity of 1500 μmol m-2 s-1, and with a CO2 concentration of 400 μmol mol-1 with a constant flow rate of 500 μmol s-1. The measured variables were the maximum photosynthesis (Amáx) (μmol CO2 m2 s-1), transpiration (E) (μmol H2O m2 s-1) and stomatal conductance (g) (mmol m2 s-1). The efficiency of water use was determined during the phenophases of heading, flowering and filling of the grains by means of the quotient of photosynthetic activity and transpiration (Zhang et al., 2016).

Agronomic evaluation

In the plots, we evaluated the plants height (AP) (cm), spike length (LP) (cm), mass of the spike (MP) (g), number of full (GLL / P) and vain grains (GVP) by ear (number), mass of the grains (MG) (g), mass of a thousand grains (M1000) (g) and yield (REND) (kg ha-1). The height of the plant was measured in 40 plants with a measuring tape (TRUPER B122080) of 3.0 m, and from the base of the stem to the apical end of the spike.

The length of the spike was measured in 30 spikes per plot, taken at random, from the base to the terminal grain with a millimeter ruler (PILOT) of 50 cm. The mass of the spike was determined in 40 spikes per plot using a SUIM LAB technical scale with an error of 0.0012 g. The number of full and vain grains per spike was quantified in 25 spikes taken at random, each spike was carefully shredded and the number of full and vain grains counted.

The grain mass was measured in 200 grains taken at random, with a technical balance (Sartorius CP64-OCE) (error of 0.001 g). The mass of a thousand grains was measured in 10 groups of a thousand grains, in the manner used for the mass of the grains. The yield for each plot was determined based on areas of 1 m2, the total mass of grains was quantified in each m2 of surface, and the yield was calculated by dividing the total mass and the area of the plot.

Experimental design and statistical analysis

Experimental design

For the genetic stability index, the experimental design was completely randomized, taking 10 representative sites from each valley, and from each site six repetitions were taken.

Data processing

With the variables AP, LP, GLL and REND evaluated in eight years and the two valleys we estimated the genetic and environmental variances, the heritability, an index of adaptability and general stability (Wi), as well as the indices for favorable environments (WiF ) and for unfavorable environments (WiD), according to the methodology proposed by Annichiarico (1992). The comparison of means of the determined indices was carried out through the GENES program (Cruz, 2013).

The data of the components of agricultural yield to determine the genetic stability index in the period between the 2007-08 to 2014-15 crop cycles were compared by means of a double ANOVA classification based on a random-effect linear model; and the components of variance were separated in the analysis of both factors (valleys: Valles del Yaqui and del Mayo) and crop cycles: (2007-08 - 2014-15). The averages were compared with the Tukey test (p ≤ 0.05).

For the statistical processing of the variables evaluated in CETT-910 we used the theoretical distribution of probabilities t of Student with the comparison of the value of each variable evaluated with the value of the varietal descriptor, an unknown population standard deviation and a sample size of n <30.

To establish the differences in photosynthetic activity, transpiration and the efficiency of water use in leaves and spikes (without interaction between organs) we compared the means between phenophases with the Tukey test (p ≤ 0.05) after performing the ANOVA of simple classification (without organ-phenophase interaction). In this variable, the unadjusted determination coefficient (R2) was determined to analyze the contribution percentage of the phenophases in the response variability obtained.

For the NDVI measurements, a phenofasic curve was constructed, taking the NDVI value as the ordinate axis, and the phenophases as abscissa. In each ordered pair, the standard deviation of the mean was observed. For these statistical analyzes we used the STATISTICA program, version 8.1, (StatSoft, 2008) for WINDOWS.

Results and Discussion

Genetic stability index of the CIRNO C2008 variety

Wheat, like other crops, fluctuates within a single crop between years and different sites due to the polygenic nature of grain yield components, and these variations depend on the ability of the genotype to adapt to changing conditions such as availability of water and nutrients, solar radiation, temperature, and other factors (Shirdelmoghanloo et al., 2016).

The results showed a significant interaction between the years (cultivation seasons) and the valleys, with the exception of the GLL variable. The variables AP and REND showed low values, and moderate LP and GLL, which denotes the little dispersion and considerable homogeneity of the data obtained for these characteristics, as an indirect indicator that the variety CIRNO C2008 did not have significant variations in its agronomic performance during the years and the valleys where it was evaluated (Table 1).

Table 1 Mean squares of ANOVA applied in four characteristics [plant height (AP), length spike (LP), filled grains (GLL) and yield (REND)] in eight seasons and two locations for the CIRNO C2008 variety. 

Fuentes de variación Cuadrados medios
gl AP LP GLL REND
Valles 1 0.53 ns 3.72 ns 0.001 ns 1.99 ns
Épocas 7 0.71 ns 11.91 ns 81.60 ns 3.04 ns
Valles x Épocas 7 0.63 14.16 475.49 ns 0.85
Residuo 48 0.11 0.69 296.65 0.01
Media 89.55 3.70 55.64 6.10
C. V. (%) 0.37 22.50 30.95 1.11
h2 89.12 73.15 57.00 49.76
Correlaciones de Pearson LP 0.19 - - -
GLL -0.19 0.07 - -
REND -0.05 -0.14 0.35 -

ns Not significant (p> 0.01) for the test of F. CV: coefficient of variation; h2: heritability. gl: degrees of freedom.

The four characteristics evaluated had low correlations between them and only r (REND - GILL) = 0.35 was statistically different (p ≤ 0.01). The highest values of heritability were found in the variables plant height and spike length, indicating the lower negative incidence of environmental factors. The number of filled grains and grain yield showed heritability values close to 50%, which indicates greater sensitivity to changes in the environment.

Obtaining a coefficient of low genetic variation for AP, REND and LP in different localities and eight years denotes the presence of monogenic dominant characters (Gouache et al., 2016) or the superdominance of minor genes in the groups of genes that affect positively the grain yield (Mirabella et al., 2016).

The polygenic character of the grain yield of cereals in general, and of wheat in particular, is the main limitation for genetic improvement programs, but numerous genes have been studied that overexpress and show superdominance, although they have been genotypes exposed to the environment for a long time. Many of these genes or groups of genes were characterized and used for the differential selection of genotypes among diverse germplasms or between parents in genetic crosses (Vázquez et al., 2016).

The genetic stability of morphological (Liu et al., 2016) and agronomic characters is important because they can be useful in any breeding program or for the use of genotypes as a model to simulate or predict possible responses to stress conditions (Vázquez et al., 2016).

The negative indices (Table 2) represent unfavorable times for the evaluated variables. The variables GLL and REND presented similar classification as favorable or unfavorable at the same times. The classification result was similar for the other two variables, with the exception of the first period (2007/08).

Table 2 Genetic stability indices (IEG), average of plant height (X-): (AP), length of spike (LP), filled grains (GLL), grain yield (REND) and classification in favorable (F) and unfavorable (D) environments of two valleys in eight evaluated seasons (p ≤ 0.05). 

Épocas AP (cm) LP (cm) GLL (unidad) REND (kg ha-1)
X- IEG Clasif. X- IEG Clasif. X- IEG Clasif. X- IEG Clasif.
1: 2007/08 89.68 0.13 F 3.19 -0.51 D 49.00 -6.64 D 5.80 -0.30 D
2: 20080/9 89.18 -0.38 D 1.94 -1.76 D 57.80 2.16 F 6.33 0.23 F
3: 2009/10 89.33 -0.23 D 3.66 -0.04 D 57.39 1.75 F 6.23 0.13 F
4: 2010/11 89.85 0.30 F 5.53 1.82 F 59.38 3.73 F 6.99 0.89 F
5: 2011/12 89.40 -0.15 D 2.96 -0.74 D 57.40 1.76 F 6.85 0.75 F
6: 2012/13 90.01 0.46 F 3.94 0.24 F 54.88 -0.77 D 5.90 -0.20 D
7: 2013/14 89.29 -0.36 D 3.04 -0.66 D 54.64 -1.00 D 5.36 -0.74 D
8: 2014/15 89.68 0.13 F 5.34 1.64 F 54.66 -0.98 D 5.34 -0.76 D

The 2011/12 season maintained the best environment for all the performance components evaluated in both valleys, classified as favorable, while 2013/14 had an opposite effect.

According to the indices described by Annichiarico (1992) (Table 3), (Wi) the Valle del Mayo showed better results than the Valle del Yaqui regarding the variables evaluated, with the exception of REND whose higher values occurred in the Valle del Yaqui. Similarly, in this valley the indices for favorable environments (WiF) were higher for AP and GLL.

Table 3 Indices of general stability (Wi), index for favorable environments (WiF), index for unfavorable environments (WiD) and average of four characteristics for each index when evaluating the Yaqui and Mayo valleys in eight cultivation seasons (p ≤ 0.05). 

Valles Altura de la planta (cm)
Media general Wi Media en AF WiF Media en AD WiD
Valle del Yaqui 89.48 99.40 89.92 99.90 89.04 99.21
Valle del Mayo 89.62 99.57 89.69 99.64 89.56 99.79
Longitud de las espigas (cm)
Media general Wi Media en AF WiF Media en AD WiD
Valle del Yaqui 4.74 39.59 4.48 12.39 4.83 43.97
Valle del Mayo 6.81 69.34 6.38 62.43 6.54 67.54
Granos llenos (unidad)
Media general Wi Media en AF WiF Media en AD WiD
Valle del Yaqui 55.53 89.95 58.86 100.55 52.19 83.94
Valle del Mayo 55.76 90.84 57.12 97.53 54.39 88.73
Rendimiento (kg ha-1)
Media general Wi Media en AF WiF Media en AD WiD
Valle del Yaqui 6260 92.94 6730 99.76 5790 88.83
Valle del Mayo 5940 87.40 6460 95.77 5410 81.76

AF and AD are averages for favorable and unfavorable environments, respectively.

In the present analysis (Tables 2 and 3) the classification is more associated with the variation of the valleys than with the different seasons. The indices obtained, with the exception of the spike length (LP) that had a greater difference between the values of Wi, WiF and WiD, did not register statistical differences in the other three variables, which indicates little variation or high stability of the morphological and agronomic characters of the cultivar in both valleys.

The Wi index was higher in the Valle del Mayo, except for the REND characteristic that had the highest values in the Valle del Yaqui. This type of genotype stability analysis is used in the selection of wheat cultivars (Benin et al., 2014), to identify genotypes of predictable behavior that respond to environmental variations under specific (WiF and WiD) or general (Wi) conditions.

Physiological and agronomic characteristics of the variety CIRNO C2008 in the CETT 910 in the 2015-2016 crop cycle

The phenology of the variety CIRNO C2008 had no significant changes due to the effect of the cultivation area and there was only a significant difference with the values of the varietal descriptor (Figueroa et al., 2010) in the phenophase of heading. This variation did not influence the subsequent phenophases (Table 4).

Table 4 Main phenological stages of the CIRNO C2008 hard wheat variety in the CETT-910 of the Valle del Yaqui. Varietal descriptor (Figueroa et al., 2010). 

Variedad Principales fenofases (días).
Macollamiento 1er nudo Espigamiento Llenado del grano Maduración
CIRNO C2008 28 ns 36 ns 75** 84 ns 120 ns
Descriptor 28±1 36±1 80±1 85±1 120±2

** p≤ 0.01; ns: not significant (t-Student test; p > 0.05).

Commercial varieties can shorten or lengthen the time of a phenophase due to the edaphoclimatic conditions in the region where they are established (Kurepin et al., 2015). The most frequent in studies of exposure and tolerance to stress conditions is a reduction of phenophase time (Guzmán et al., 2016), which accelerates senescence and abscission, especially during physiological drought associated with drought or salinity stress (Mendes et al., 2016).

For thermal stress conditions, the time of phenophases decreases due to the increase in the concentration of gibberellic acid (Argentel et al., 2016). The time of the crop phenophases is an efficient indicator of the good vegetative and nutritional state of the plants, and has a positive correlation with the yield, but it is necessary to carry out complementary studies such as the vegetable biomass index in each phenophase (Ballesteros et al., 2016).

Normalized difference vegetation index

The NDVI evaluated in the phenophases of the CIRNO C2008 crop had a significant increase since the tillering phenophase and the maximum point was obtained in the heading (Figure 2). This result shows the good physiological and nutritive state of the plant. The decrease in the NDVI values since heading is probably due to the translocation of nitrogen compounds, which favors the filling of the grains and is an important sign of the senescence of the plants (Gaju et al., 2016).

Figure 2 Normalized difference vegetation index in the main phenophases of the CIRNO C2008 wheat variety in the CETT during the 2015-16 crop cycle. The bars represent the standard deviations of the measurements in each phenophase. 

Measurements of the nutrient status of plants in the different phenophases of the crop through the Greenseeker has been very successful in the intensive and extensive wheat systems and allows to quantify the efficiency of the use of nitrogen fertilizer, as well as to predict the grain yield (Stefen et al., 2016).

Photosynthesis, stomatal conductance and transpiration

The photosynthetic activity in the phenophases of heading, flowering and grain filling was high in the leaf (Table 5), but the highest value was obtained at the heading, although the values of stomatal conductance and transpiration did not present significant differences in the phenophases. The decrease of the photosynthesis values after heading may be due to the function of the accelerated mobilization of photoassimilates from the sources (leaves) to the drains (grains) (Shirdelmoghanloo et al., 2016).

Table 5 Photosynthetic activity, stomatal conductance and transpiration of the CIRNO C2008 variety in the phenophases of heading, flowering and grain filling. 

Fenofases Fotosíntesis (A). Transpiración (E) y Conductancia estomática (g)
A(µmol CO2 m2 s-1) E (µmol H2O m2 s-1) g (mmol m2 s-1)
hoja espiga hoja espiga hoja espiga
Espigamiento 25.3±1.2 a 16.4±2.3a 4.5±0.03 ns 1.2±0.01 ns 293.4±14.6 ns 128.1±8.3 ns
Floración 23.8±1.4 b 13.5±1.2 b 4.6±0.07 ns 1.4±0.02 ns 294.6±11.4 ns 126.4±5.2 ns
Llenado de granos 22.6±2.3 c 2.7±0.5 c 4.4±0.01 ns 1.4±0.01 ns 293.1±13.2 ns 117.9±8.6 ns
R2 0.99 0.98 0.99 0.97 0.98 0.99
CV 3.17 2.46 0.1 0.1 0.26 4.06

Means with different letters in a column are statistically different (Tukey, p ≤ 0.05).

ns: not significant.

The photosynthetic activity in the spikes was lower than in the leaves in the three phenophases evaluated (Table 5). These values decreased when these phenophases passed; however, the values were greater than 10 μmol CO2 m-2 s-1 in the phenophases of heading and flowering, which represents a contribution of this organ to the grain yield. The photosynthesis in the spikes showed a highly significant decrease during the filling phenophase with respect to that obtained in heading and flowering.

Transpiration and stomatal conductance in the spikes were lower than in leaves and there were no differences between the phenophases. Photosynthesis in the spikes contributes to a good filling of the grains although there are divergences with respect to the percentages of contribution, due to the form of evaluation of the contributions and the variety (Borrás et al., 2004).

Contributions (although in small amounts) of the photosynthetic activity of organs with a lesser degree of specialization, have a biochemical and agronomic importance because the photoassimilates of the leaves are destined to the fruit and to all the organs of the plant to maintain metabolism (Zhang et al., 2016).

Obtaining during the phenophases of flowering and grain filling high photosynthesis values in the leaves (Table 5), higher than 20 μmol CO2 m-2 s-1 (Saeed et al., 2017) and a stable transpiration, denotes the capacity of the variety to ensure a good grain yield. A well-nourished plant, with adequate moisture in the soil and sufficient solar radiation, maintains a considerable photosynthetic activity so that a greater grain yield can be expected (Carmo-Silva et al., 2015).

The photosynthetic activity is the main physiological variable that contributes to the correct filling of the grains, their final mass and yield. When photosynthesis is low in reproductive phenophases there is miscarriage of the distal spikelets or poor filling of the grains, which leads to significant losses in grain yield and quality (Prins et al., 2016).

There are not enough references about the measurement of the photosynthetic activity and phenophases depending on the NDVI in the variety of wheat studied, therefore the results obtained will contribute to the physiological characterization. These results could be used in breeding programs, since the photosynthetic activity defines the progress in wheat genetic improvement (Vázquez, et al., 2016).

Water use efficiency (WUE) based on the photosynthesis-transpiration ratio

The WUE was similar in the leaf in the three phenophases evaluated (Figure 3); however, in the spikes, although it was higher in the phenophases of heading and flowering than in the leaf, there was a significant decrease in their values after the phenology, which could be 99 % due to the effect of the phenophases.

Figure 3 Water use efficiency in leaves and spikes of the CIRNO C2008 variety during the phenophases of heading, flowering and filling of the grains in the CETT in the 2015-16 cycle. R2: coefficient of determination, without adjustment, of the WUE variation in the different phenophases. 

The decrease in the photosynthetic activity of the spikes was the factor that most influenced the WUE variations (Table 5), which is attributed to the loss of the chlorophyll pigments as the spike grows (Jamil et al., 2016), the rupture of chloroplasts due to the concentration of ethylene during maturation, and the increase in the activity of the chlorophyllase enzyme that accelerates chlorophyll catabolism (Sánchez et al., 2016).

The highest value of WUE in the spikes was obtained during the heading, which is due to the fact that the spikes have, in their green state, considerable advantage for capturing the light with respect to the flag leaf and the remaining leaves, as a result of the little or no interference from shading or high irradiance, allowing it to maximize photosynthesis (Blum, 2005).

In addition, low transpiration also contributed since the spikes are not bodies specialized in the realization of this process. This response was the main contributor to WUE superiority of spikes over the leaf in the present study.

The evaluation of WUE determined by the photosynthesis/transpiration ratio is an important indicator to select genotypes with high photosynthetic capacity in conditions of low water availability, and tolerance to drought stress is attributed to it (Messina et al., 2015).

These genotypes might be tolerant to thermal stress, although it is assumed that this type of stress considerably increases transpiration to achieve thermoregulation of the leaf mesophyll (Cowie et al., 2016).

Grain yield components

Decreases in the spike mass (5.35 %) and the number of full kernels (3.3 %) were significant with respect to the varietal descriptor (Table 6), although there was no negative correlation between these variables and agricultural yield (r (MP- REND) = 0.012; r (GLLP-REND) = 0.018). On the contrary, the dry mass of the grain and the grain yield were statistically higher.

Table 6 Measurement of the components of the agricultural yield of the hard wheat CIRNO C2008 variety evaluated in the CETT- 910 of the Valle del Yaqui. 

Componentes del rendimiento
Fuente de variación AP LP MP GLL P-1 M1000 Masa seca REND
paja grano
CIRNO C2008 88±2ns 7.1±0.1ns 53±1.1 58±1 60.6±0.1ns 3.43 0.7 7400±0.2
Descriptor 78- 90 7.0±2 56 ±1.1 60±1 60.5±1 4.0 0.58 6400±0.2

ns Not significant (t-Student test). AP: height of the plants (cm); LP: length of the spike, (cm); MP: spike mass (g); GLL P-1: number of filled grains per spike. MMG: mass of a thousand grains (g). Dry mass of straw and grain: (kg ha-1). REND: yield (kg ha-1).

Obtaining a superior yield in relation to the varietal descriptor shows that the variety still maintains its genetic-productive potential, perhaps due to the ability of the plant to mobilize the photoassimilates from the foliage to the spike and propitiate an adequate filling of the grains. (Vázquez et al., 2016). This can be corroborated by the significant difference found between the straw and grain dry mass values. The dry mass relation in terms of grain/straw can vary because the edaphoclimatic conditions and of cultivation affect the translocation and distribution of photoassimilates from the sources and reservoirs up to the sinks (Hortelano et al., 2013).

The variables plant height and spike length did not present significant differences with respect to the varietal descriptor (Table 6), which shows the genetic stability of the variety during its exposure to field conditions. This is notable because most of the yield components have a polygenic nature but perhaps there are minor genes that still maintain their dominant character before a considerable variability of edaphoclimatic conditions (Asseng et al., 2015). In wheat, plant height and in particular the NDVI are efficient variables for the estimation of yield, and are reported of being precise indicators for differential selection programs (Pantazi et al., 2016).

Conclusions

The CIRNO C2008 wheat variety maintains its genetic stability after eight years of release for agricultural production in the Valles del Yaqui and del Mayo, based on the variables evaluated. The variables plant height and spike length were the components of the yield with greater genetic stability. The highest grain yield was obtained in Valle del Yaqui

The phenology of the variety does not experience significant variations with respect to the varietal descriptor after eight years of being released for agricultural production in Sonora and, in addition, maintains its grain yield close to its productive genetic potential, high levels of photosynthetic activity and water use efficiency.

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Received: January 2017; Accepted: October 2017

*Autor para correspondencia: garatuza@itson.edu.mx

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