GHS101 and GHSP26 improves boll retention in upland cotton grown under high temperature of semi-arid subtropical climate

doi:10.21203/rs.3.rs-1643410/v1

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GHS101 and GHSP26 improves boll retention in upland cotton grown under high temperature of semi-arid subtropical climate

https://doi.org/10.21203/rs.3.rs-1643410/v1

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Background

Cotton is a natural fibre producing crop grown worldwide where agro climatic conditions match with its growth and reproduction. High temperature is a major environmental factor affecting cotton production due to climate change. Only way to cope with high temperature is to breed for the tolerance. High temperature tolerance is a complex mechanism that includes many genes by which cotton responds to stress. Genes conferring heat stress can be best studied in segregating population like F₂ grown under stressed environment.

Results

Based on contrasting traits (boll retention percentage), two cotton cultivars i.e. FH-Lalazar (High temperature susceptible) and CRIS-9 (High temperature tolerant) were evaluated for eight heat stress related genes. Out of eight, two genes (i.e. GHS101 and GHSP26) were present only in tolerant genotype i.e. CRIS-9 and these two were selected for further analyses. After development of F₂, the morphological and yield data from parental as well as F₂ population was recorded under natural high temperature conditions. The data for boll retention percentage under high temperature was recorded as an indicator of high temperature tolerance. These two genes were surveyed in 129 plants of F₂ population. Most of the plants, containing both genes showed up to 81% boll retention, but some exceptions were there. Most of the phenotypic traits showed skewed variation on histogram depicting metric nature of genes.

Conclusion

Results demonstrated that co-segregation of boll retention property along with heat tolerant genes in F₂ population is important in reproductive development of cotton under high temperature conditions. Thus, it is necessary to identify, and pyramid other genes related to high temperature tolerance at reproductive stage.

Boll retention

Gossypium hirsutum

Climate change

Heat shock protein HSPs

Climate change is changing paradigm of field conditions by unpredictable heatwaves, rainfall, and humidity. Major crops are considered vulnerable to yield losses by heat and drought stress due to adverse climate change (Vinocur and Altman 2005; Iqbal et al. 2018). Cotton (Gossypium hirsutum L.) originated from tropical and subtropical regions, considered to be better inhabited in hot and semi-arid environments (Singh et al. 2007) but it appeared that heat and drought stress have drastic effect on productivity and quality (Lokhande and Reddy 2014; Iqbal et al. 2019). Photosynthesis decreases significantly when temperature reaches above 36 ^oC. Increase in temperature affects shoot development, flower/bud formation, bolls/plant and quality of fiber (Khan et al. 2014; Farooq et al. 2015).

In Pakistan where temperature reaches to 50°C during May and June, which is 20 ^oC above than the optimum temperature (Burke et al. 2004; Mohamed and Abdel-Hamid 2013), due to which cotton crop faces severe heat stress. Cultivar development against heat stress struggles for potential breeding material, as cultivated germplasm showed low variation (Van Esbroeck 1999; Rungis et al. 2005), maybe due to using same material in breeding programs (Wendel et al. 2009; Iqbal et al. 2019), causing future breeding for stress tolerance very difficult. Secondly, conventional breeding methods requires a lot of time (Dabbert and Gore 2014) and resources where scientists required to delay for selection till crop maturity. Conventional breeding along with molecular breeding is the solution for development of potential cultivars against abiotic stress. Molecular analysis for abiotic stress tolerance (heat, drought, salinity) have identified specific genes controlling the abiotic stress tolerance and are used by many scientists for screening genotypes for abiotic stress tolerance (Rodriguez-Uribe et al. 2014; Chen et al. 2015; Demirel et al. 2016; Iqbal et al. 2017).

High temperature affects vegetative as well as reproductive growth of cotton but reproductive growth is more prone to it. At reproductive stage, high temperature cause drop-age of buds, reducing bolls/plant and ultimately lead to reduction in yield (Zhao et al. 2005; Snider et al. 2010; Iqbal et al. 2017). Plants may tolerate high temperature (Willits and Peet, 1998) by adopting some physiological changes. Some plants have ability to adopt high temperature environment due to the presence of genes which regulate growth and development under high temperature. Many genes are identified as stress responsive genes. Among these, heat shock proteins (HSPs), produced in response to sudden change in temperature, are considered as the most important one (Hasanuzzaman et al. 2013). Photosynthesis (Salvucci et al. 2004; Snider et al. 2010; Xiao et al. 2017) and cell membrane stability (Levitt 1980) is the first target of high temperature in response heat shock proteins helps normal photosynthesis and maintains integrity of cell membrane (Ristic and Momcilovic 2007; Mohamed and Abdel-Hamid 2013). These proteins are produced immediately after temperature exceeds from optimum level (Hasanuzzaman et al. 2013) and helps improve the physiological response in cotton against heat stress (Ristic and Momcilovic 2007). Zhang et al. (2005) enlisted few important heat tolerant genes in cotton that help improvement in cotton response against high temperature. The selected genes GhPP2A1, HSFA2, GHSP26, HSP3, HSC70-1, HSP101, ANNAT8 and APX1 were screened in cotton genotypes to identify tolerant one (Zhang et al. 2005). These genes are well reported for their role in growth and development of cotton under heat and drought stress. But their role in reproductive development, especially boll retention under the stressed environment is still not clear.

Iqbal et al. (2017) reported that even with presence of heat tolerant genes, some genotype produce less yield and they found that such genotypes performed well for vegetative growth and suggested to correlate reproductive traits with heat tolerant genes for efficient marker assisted section. Boll retention property of cotton under high temperature is very important for stability of yield in un-predictable environments. Genetics of the trait and genes associated with boll retention has not been extensively studied. Segregation study of the trait may provide knowledge of genetic basis of tolerance against high temperature in cotton. The present study has been conducted to assess association of boll retention and heat tolerant genes in segregating population developed from a cross between two diverse cotton genotypes with the aim of development of heat tolerant advance lines.

Experimental site and Plant material

The experiment was conducted at research farm of Bahauddin Zakariya University, Multan weather is characterized as subtropical semi-arid with hot summer and cold winters. In summer season (late May and June) temperature often crosses 50 °C. Therefore, cotton crop grown in April and early May face high temperature at the start of reproductive stage. In this experiment, we applied natural high temperature by sowing the crop in early April to have maximum implication in the field. Two parents [FH-Lalazar (High temperature susceptible) and CRIS-9 (high temperature tolerant)] were selected from genotypes based on morphological data as well as boll retention percentage under high temperature. The variety FH-Lalazar was developed at Cotton Research Station Faisalabad whereas CRIS-9 was developed at Cotton Research Institute Sakrand, Pakistan. True to type seed of the varieties was collected from the respective research institute to rule out any seed adulteration. The selected parents were grown in the glasshouse under controlled conditions. The parents were crossed following standard protocol to produce F₁. At maturity, F₁ seed was harvested from plants and sown in the field. To produce F₂ seed, flowers of F₁ plants were bagged to avoid contamination and tagged for reference. The segregating F₂ population was developed and grown in the field in first week of May 2019, following all cultural practices. Reason for early May sowing was that maximum of the flowering can occur at peak temperature to apply natural high temperature (Iqbal et al. 2019).

Data collection

The morphological data were recorded from a total of 129 plants of F₂. For other morphological and yield related data, individual plants were screened for final plant height (cm), counts of monopodial and counts of sympodial branches, total counts of bolls per plant, seed index, counts of nodes to first monopodia as well as ginning out turn (GOT%).

Data for boll retention percentage was collected throughout the season where the reproductive phase started from mid-June, when the average temperature was 40⁺ °C and flowering retain or shedding contributed to the total boll retention. Boll retention percentage was calculated as by (Iqbal et al. 2017).

Fibre quality analysis was conducted through HVI machine available at Central Cotton Research Institute Multan, for staple length (mm), fibre strength (g/tex) and fibre fineness (U).

Molecular studies

Eight selected genes, GhPP2A1, HSFA2, GHSP26, HSP3, HSC70-1, HSP101, ANNAT8, and APX1 (Additional file 1: Table A1) to select the contrasting genes. Two genes (GHS101 and GHSP26) were further selected for molecular characterization the F₂ population for boll retention percentage. Primer sequence of all the genes are presented in Additional file 1: Table A1.

DNA was extracted from parental (P₁ and P₂) population as well as segregating (F₂) population as by (Doyle and Doyle, 1987). Selected gene segments were amplified through PCR for about 35 cycles for each of the gene specific primers used in the reaction mixture. Reaction mixture consisted of the resulted bands were examined in gel documentation system using agarose gel (1.5 %) and the fluorescent dye used as ethidium bromide. Scoring of the bands was done based on the desired gene presence or absence. The genes which were present in heat tolerant parent and absent in heat susceptible parent were selected to screen F₂ population in Additional file 2: Table 2.

Statistical analyses

All collected data were undergone statistical analyses. Data for all traits of F₂ population was used to develop histogram to understand the nature of gene controlling them. F₂ data were further analysed for correlation analyses as developed by (Dewey and Lu, 1959).

Inheritance nature of the selected genes was analysed by means of Chi-square analysis according to (Pearson 1899) as per following formula.

$${{\chi }}^{2}=\sum \frac{{(\text{O} - \text{E})}^{2}}{E}$$

Where: O is the observed frequency of specific gene and E is the expected frequency of specific gene as per Mendel’s law of segregation.

Morphological, yield and fiber quality

All morphological and yield (Except number of locules and monopodial branches) traits showed continuous variation in the segregating population. Plant height range between 60 cm to 170 cm where 103 plants were medium heighted 90–100 cm. Range of sympodial branches counts per plant was 4–24. Total counts of bolls/plant varied from 10–45. The range of seed cotton yield was 50-150g, indicating continuous variation in important yield traits. Average boll weight per plant was from 0.5-2.5g under high temperature. The trait boll retention percentage, taken as key parameter for assessment of heat tolerance in cotton under high temperature, also shows continuous variation in F₂ population. Boll retention ranged 45–82% in 129 plant of F₂ population (Additional file 3: Table. A3). Lint weight per plant ranged between 20-90g showing eight classes in F₂ population. Total number of locules showed minimum variation, only 4 or 5 locks/boll were observed. Monopodial branches ranged from 0–1 whereas three plants had two monopodial branches (Fig. 1). Thus from variation in F₂ population, these traits seem to be controlled by one gene. Fiber traits like, ginning out turn, fiber strength, fiber length and fibre fineness also showed medium amount of variation (Fig. 1).

Molecular Characterization
The parents FH-Lalazar and CRIS-9 have been screened for eight genes GhPP2A1, HSFA2, GHSP26, HSP3, HSC70-1, HSP101, ANNAT8 and APX1 (Fig. 2) and observed that heat susceptible variety FH-Lalazar (boll retention = 57%) lacks two heat shock protein genes GHSP26 and HSP101 (Additional file 2: Table A2), while heat tolerant variety CRIS-9 (boll retention = 84%) appeared possessing all eight genes. The gene GHSP26 appeared in band of about 213 kb in size on gel and the gene HSP101 showed band size of 200 kb. The F₂ population (129 plants) were screened for these two genes using gene specific marker (Fig. 3). GHSP26 specific marker amplified in 45 plants and HSP101 amplified in 43 plants. It is of interest that all the plants having both genes showed 71% or more boll retention. Similarly plants having anyone of the gene showed at 61% or more boll retention. Moreover, any plant having none of the genes from these two, showed 67% or less up to 45% boll retention (Additional file 3: Table A 3).

Heat tolerance in cotton being polygenic trait, is difficult to manipulate in breeding program. Falconer and Mackay (1996) described that the simultaneous segregation of many genes controlling the trait shows continuous variation. The traits under study appeared as continuous variation on frequency distribution graphs (Fig. 1). The trait boll retention percentage showed continuous variation on histogram, indicating it a polygenic trait. The trait is considered as a key indicator of high temperature tolerance in cotton (Zhao and Oosterhuis 2002; Zhao et al. 2005; Snider et al. 2010; Iqbal et al. 2017). Correlation results also explains the importance of boll retention for higher yield of cotton (Additional file 4: Table A4).

In this experiment we have used boll retention as a key criterion for selection against high temperature. In this experiment high temperature was applied by early sowing of cotton which results in high temperature on early reproductive development where temperature crossed 40°C after 45 days of cotton sowing when reproductive growth normally starts. In high temperature susceptible plants, most of early developed buds dropped which results in lower final seed cotton yield. Detrimental effect of high temperature on reproductive development of cotton already have been reported in literature (Snider et al. 2010; Iqbal et al. 2018; Singh et al. 2018). Correlation analyses strengthen this hypothesis where boll retention is strongly correlated not only with yield but also with other yield contributing parameters like counts of sympodial branches, plant height and lint weight. Boll retention showed non-significant (p ≥ 0.05) negative correlation with boll weight which depicts that with increasing number of bolls, relative boll weight decreased (Munir et al. 2016; Iqbal et al. 2018), but as boll weight is also a genetically controlled traits therefore, this effect is little one.

GHSP26 and HSP101 are well known for their role in high temperature tolerance (Myouga et al. 2006; Maqbool et al. 2010; Zafar et al. 2018). But it is not clear that whether these genes are associated with reproductive growth or these only promote vegetative growth under high temperature. For example, eight genes for high temperature tolerance were screened for the parents, out of which six were present in heat susceptible parent (based on reproductive traits), but they promoted only vegetative growth. Boll retention is highly affected by high temperature offered through early sowing of cotton (Iqbal et al. 2017). In current study, boll retention ranged from 45–82% in F₂ population. Data for each plant for boll retention and gene amplification was collected separately (Additional file 3: Table A3). The co-segregant analysis indicates that in F₂ population, the higher percentage of boll retention percentage appeared associate with the presence of both genes GHSP26 and HSP101 in Additional file 3: Table A3.

The results have shown that these two genes are associated with high boll retention. Plants where both genes were presented showed up to 82% boll retention while plant with only one gene of the two showed up to 71% boll retention with some exception even higher. It is worthy note-able that even with the presence of one gene of the two minimum boll retention was 62% while maximum boll retention without any gene was recorded up to 67%. This strengthen the reality that high temperature is a polygenic trait and is controlled by many gene (Singh et al. 2007; Jha et al. 2014; Iqbal et al. 2017). But as most of the plants having both genes showed maximum high temperature tolerance, this showed that many heat tolerant genes associated with reproductive development of cotton are linked with these two genes. This may be because of the dropping of flowers/buds is a physiological phenomenon in cotton, and two genes helps improvement in physiology of cotton plant under sudden high temperature. These genes are considered very important for heat tolerance in cotton as they produced heat shock proteins (HSPs) in response to rise in temperature from optimum level. The proteins help maintains stability of cell membrane and normalize process of photosynthesis (Ristic and Momcilovic 2007; Mohamed and Abdel-Hamid 2013). Further physiological processes may be identified about involvement of these genes for flower/bud retention in the cotton under high temperature.

Chi-squire test for supported the 3:1 hypothesis for both genes GHSP26 and HSP101 in F₂ population. The gene GHSP26 appeared in 45/127 plants of F₂, whereas the gene HSP101 appeared in 43/127 plants of F₂ and thus confirmed the recessive behaviour of these two genes.

As it is revealed from the Chi Square analysis that the both genes fit well into Mendel’s law of segregation and are inherited as recessive alleles, therefore in successive generations, more plants with presence of the genes may also appeared. Development of cotton genotypes with pyramiding of these both genes may produce higher yields in climate change scenario.

GHSP26 and HSP101 genes are well associated with boll retention under high temperature environment in cotton. As these genes are recessive in nature therefore, selection may be delayed in successive generations to find out maximum combinations. Development of cotton genotypes with pyramiding of these both genes may produce higher yields in climate change scenario. As high temperature tolerance is a polygenic phenomenon, more genes may be identified associated reproductive development under high temperature to produce maximum cotton yields under climate change scenario.

Ethics approval and consent to participate

Not applicable.

Consent for publication

Not applicable.

Availability of data and materials

The data sets used and/or analysed during the current study are available from the corresponding author on reasonable request.

Competing interests

Authors declare no conflict of interest for this study.

Funding

The study was part of National Research Projects for Universities (NRPU) by Higher Education Commission Pakistan, entitled “Marker assisted gene pyramiding for heat tolerance in cotton” Project ID: 7965. We are thankful to HEC for providing funds for this research work.

Author contribution

Research supervisor, Muhammad AS; Co-supervisor, Waqas M; Manuscript, Muhammad A; Lab work, Shakeel A; Data collection, Mirza MAB; Analysis, Muhammad WA; Lab supervision, Muhammad QA; Review and editing, Hafiz MWA.

Acknowledgment

We would like to thank the all reviewers and authors for their valuable comments and helpful suggestions which help to improve the manuscript. Meanwhile, I am thankful to Dr. Muhammad Asif Saleem giving help for this paper.

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GHS101 and GHSP26 improves boll retention in upland cotton grown under high temperature of semi-arid subtropical climate

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Abstract

Background

Results

Conclusion

Figures

Background

Materials And Methods

Results

Morphological, yield and fiber quality

Discussion

Conclusion

Declarations

References

Supplementary Files

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