Heat Stress on Wheat Grain Yield and Quality

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1		Daniel K. Y. Tan	--	2253	2022-07-05 13:06:37	\|
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This entry is adapted from the peer-reviewed paper 10.3390/agriculture12060886

Climate change threatens to impact wheat productivity, quality and global food security. Maintaining crop productivity under abiotic stresses such as high temperature is therefore imperative to managing the nutritional needs of a growing global population.

climate change food security nutrition wheat quality global warming

1. Introduction

In the context of climate change, various abiotic stresses including rising temperatures are significantly impacting the growth and development of crops. Since 1910, atmospheric temperature in Australia has risen by an average of 1.4 ± 0.24 °C, along with an increased frequency of extreme heat events across the continent ^[1]. For instance, Australia experienced 43 extreme heat days in 2019, three times more than in any given year prior to 2000 ^[2]. Additionally, climate modelling using data published by the Intergovernmental Panel on Climate Change (IPCC) forecasts global mean temperatures to rise by between 2.0–4.9 °C by the end of this century ^[3]. Increased global mean temperatures and frequency of extreme heat days pose a significant risk to the productivity of wheat cropping systems ^[4] that rely on optimal conditions between 12.0–22.0 °C for favourable growth and development ^[5].

1.1. Wheat Grain Yield

Wheat is the third-largest cereal crop (by volume), and it is considered as one of the most important food crops for supplying the nutritional requirements of over 4 billion people ^[6]^[7]. Whilst the global population is forecast to increase to 9.8 billion by 2050, demand for food is expected to rise by 77.0% over the same period ^[8]^[9]. To adequately supply the global population in 2050, the Food and Agriculture Organisation predicts that an additional 900 million tonnes of cereals will be required to meet the demand ^[10].

Wheat cropping regions such as Australia, China and India are already experiencing high temperatures during the growing season; causing reduced grain yield ^[11]^[12]. Global wheat production is projected to decrease by up to 6.0% for each 1.0 °C increase above the optimum range of 12.0–22.0 °C ^[5]^[13]. The growth stage at which heat stress events occur has a significant impact on wheat grain yield ^[14]. Djanaguirama et al. ^[14] found that heat stress at 32.0 °C reduced yield per plant by 29.0% and 44.0% at anthesis and during the grain filling period, respectively. Due to the threat of food insecurity, there is a growing imperative to identify high-yielding wheat cultivars that display resilience to the impacts of rising temperatures and maintain nutritional and end-use quality.

1.2. Wheat Grain Quality

High temperature is well reported in the literature as among the significant factors, affecting the physical and chemical quality of wheat grains. Quality parameters, including grain number, size, grain weight, hardness and the composition of protein and carbohydrates determine the nutritional and end use value of wheat grain.

For example, grain weight and size are the important indicators of milling potential, where less flour may be extracted from smaller, more compact grains. Likewise, grain hardness is defined as the ability of grain kernels to resist mechanical force and is generally dictated by starch and protein composition within the endosperm. It is an important indicator of flour particle size and end use suitability, particularly as bread making generally requires harder wheat flour compared to biscuit and cake making ^[15].

Heat stress occurring at anthesis had a significant impact on grain number. Narayanan et al. ^[16] observed a reduction in grain number (17.0%) under a 7-d heat stress treatment at 15.0/35.0 °C, and similar results were obtained by Djanaguiraman et al. ^[14] when examining the cultivar Seri82 under heat stress at 32.0 °C. Grain weight is also susceptible to heat, particularly during post-anthesis period. Wang et al. ^[17] observed a significant reduction in mean grain weight of 20.7% across 38 diverse cultivars subjected to 37.0 °C for 3 day/night cycles at 12 days post-anthesis.

Additionally, grain protein content plays an important role in the end-use quality of wheat flour and high temperature, particularly during grain filling, and can significantly alter the quality and concentrations of grain proteins. Proteins constitute 8.0–20.0% of wheat grain, and these are divided into storage proteins (gluten) and metabolic (non-gluten) proteins. The high molecular weight gluten protein fractions gliadin and glutenin make up ~80.0% of total wheat protein ^[18]^[19]. The gliadin fraction of gluten protein provides cohesiveness and extensibility in the dough, whilst glutenins (which are further fractionalised into low and high molecular weight subunits) provide an important contribution to dough strength and elasticity ^[20]^[21]. The functionality of gluten properties, however, is dependent on the ratio of glutenin and gliadin which is found to be impacted by heat stress in the reproductive phase ^[22]^[23]. Barutcular et al. ^[24] reported a mean increase of 39.6% in protein content across all the tested genotypes at 36.3 °C, whilst multiple studies have found that heat stress can also have a significant impact on the composition of protein fractions in wheat flour ^[23]^[25]^[26].

Starch is also an important constituent of grain, comprising 60.0–75.0% of wheat yield ^[24]. The functional properties of starch are influenced by the ratio of amylose/amylopectin which can be negatively impacted by heat ^[27]^[28]. In normal starch, amylose content typically ranges between 20–30% ^[29]. A reduction in the ratio of amylose/amylopectin (from 0.22 to 0.20) was observed in the heat-tolerant cultivar Hubara-3*2/Shuha-4 when subjected to 37.0 °C temperature ^[27].

Empirical research into the effect of high temperature on wheat grain yield and quality is therefore important to securing reliable wheat yield and quality in the future climate. Identifying stable cultivars can minimise the disruption to desirable physical and chemical quality traits that govern the potential use of wheat flour in final food products. Various studies have identified a strong genotype-dependent response to post-anthesis heat in wheat. For instance, Poudel et al. ^[11] observed variation in yield between cultivars subjected to 30.0 °C; a cultivar Bhrikuti retained the highest yield of 3279 kg ha⁻¹ (a 25.5% reduction from the control at 10.0/25.0 °C (day and night temperature), whilst yield in cultivar NL 1412 was reduced by 77.1% to 754.5 kg ha⁻¹. Additionally, Hernández et al. ^[30] identified a considerable genotypic effect for protein content and gluten strength amongst 54 spring wheat cultivars under heat stress. Moreover, of 28 selected wheat lines from the International Maize and Wheat Improvement Centre (CIMMYT) in Mexico, Borlaug 100 displayed a significant tolerance to heat, producing an 8.0% higher yield compared to control cultivars Roelfs F2007 and Vorobey ^[31].

2. Classification of Wheat Cultivars

Various wheat cultivars (Triticum aestivum L. and Triticum durum L.) were identified within the current literature that displayed a range of yield tolerance to heat. For instance, a Nepalese wheat variety Bhrikuti retained the greatest mean yield (amongst 20 spring wheat genotypes) of 3279 kg ha⁻¹ under a 1-month heat treatment (max temp 35.0 °C) during anthesis, though, experiencing a 25.5% reduction from the control treatment at 10/25.0 °C ^[11]. Additionally, an Indian cultivar Raj 3765 displayed the greatest yield thermotolerance (37.5% reduction) amongst 30 different genotypes where the mean yield reduction was 59.5% with heat (35.0 °C) coinciding with the post-anthesis period ^[32]. Other significant genotypes that displayed tolerance to high temperatures included the high yielding Borlaug 100 (sourced from the CIMMYT in Mexico), and the Serbian/Bulgarian cultivar Pobeda ^[31]^[33].

Other cultivars that retained thousand kernel weight (TKW) under heat stress included the Mexican cultivar Cirno C2008 (pedigree: Sooty-9/Rascon-37//Camayo) and the Indian cultivar HD 2967 ^[23]^[28]^[32]. Cirno C2008, sown in February, retained the highest TKW of 41.6 g under heat stress at a maximum temperature between 35.0–36.0 °C during the grain filling period in May (a 21.5% reduction from the control at a maximum temperature of 32.0 °C) ^[23]. Similarly, HD 2967 retained the highest mean TKW of 30.7 g amongst 30 genotypes sown in mid-December and exposed to heat stress at 35.0 °C during post-anthesis (a 3.3% reduction from samples sown in optimum conditions in mid-November) ^[32].

Raj 4083 is a new thermotolerant wheat variety developed in Durgapura, India (pedigree: PBW 343/UP 2442//WR 258/UP 2425) that retained a TKW of 33.1 g at temperatures > 35.5 °C when sown in mid-December and exposed to post-anthesis heat, reducing by 21.4% compared to a mean TKW reduction of 31.2% across all cultivars ^[34]^[35]. Late sown Indian wheat HD 2985 retained higher TKW (23.0 g) under heat stress at 22.0/40.0 °C, 3.7 g above the mean of all genotypes compared to the control (36.6 g) at 18.0/30.0 °C, averaged across two separate growing seasons ^[36]. Moreover, PBW 621 increased in TKW by 5.9%, from 37.3 to 39.5 g at a temperature between 24.0–36.0 °C for 3 days in a controlled growth chamber ^[37]. Cultivars that could retain chemical quality under heat stress were also identified in the current literature.

3. The Effect of Heat Stress on Wheat Grain Yield

Heat Stress at Anthesis and Post-Anthesis have Varied Impacts on Wheat Grain Yield

The severity of grain yield depletion in wheat crops is highly dependent on the growth stage at which heat stress occurs. Whilst all growth stages are susceptible to some extent, heat stress during the reproductive phase (anthesis and grain filling) is particularly detrimental to grain development, which can cause reproductive sterility and significantly reduce grain number and yield ^[33]^[38]^[39].

The effect of heat stress on wheat grain yield is summarised in Table 1. Heat stress at anthesis had the greatest effect on grain number per plant. Mirosavljevic et al. ^[33] found that heat (25.0/35.0 °C) led to the cultivar Renesansa showing the highest reduction of 185 grains per plant, from 395 grains per plant at the control temperature of 16.0/24.0 °C. They concluded that the effect on grain number was likely due to flower abortion and pollen sterility at anthesis. Within the same study, the cultivar Pobeda displayed resilience to heat stress (28.0/38.0 °C) at grain filling, showing higher grain yield per plant (9 g) amongst the cultivars compared with the plants under optimum temperature (19 g) ^[32].

Table 1. The effect of heat stress on wheat grain yield at anthesis and post-anthesis.

Cultivar and Origin	Type	Heat Stress (min/max °C)	Growth Stage	Impact	Reference
38 different cultivars.	Spring Triticum aestivum L.	17.0/37.0	12 days post-anthesis.	Reduced grain filling duration by 2–9 days. The greatest reduction in grain filling duration was in the Chinese Spring variety, from a mean of 40.5 days at ambient temperature to 30 days at 37.0 °C (26.0% reduction). BJY/COC//PRL/BOW/3/Bloyka-1 had the lowest reduction from 39 days at ambient temperature to 33 days at 37.0 °C (14.8% reduction).	^[17]
Karl 92 (USA) (Hard red winter) Ventnor (Australia) (Hard white winter)	Triticum aestivum L.	15.0/35.0	7 days at anthesis.	16.0% mean reduction across both varieties. Ventnor had no significant yield reduction. Karl 92 had a significant mean reduction in yield per plant (g), decreasing from 10.0 g at optimum temperature (15.0/25.0 °C) to 6.0 g at 35.0 °C.	^[16]
Renesansa (Serbia) Pobeda (Serbia/Bulgaria) Simonida (Serbia) Gladius (Australia)	Winter Triticum aestivum L.	25.0/35.0 for all cultivars.	7 days at anthesis. Mid-grain filling 14 days post-anthesis.	The significant mean reduction in grain yield per plant for all cultivars. Pobeda retained the highest mean grain yield per plant under all treatments; 19.0 g at the control temperature (16.0/24.0 °C), 11.0 g with heat stress at anthesis, and 9.0 g with heat stress during post-anthesis. The lowest mean grain yield per plant was observed in Renesansa; 11.0 g at the control temperature, 5.0 g with heat stress at anthesis, and 5.0 g with heat stress during post-anthesis.	^[33]
	Winter Triticum aestivum L.	28.0/38.0 for all cultivars.			^[33]
Bhrikuti (Nepal) Gautum (Nepal) 3 × Bhairahawa lines (Nepal) 15 × Nepal lines (Nepal)	Spring Triticum aestivum L.	15.0/30.0	At grain filling.	47.6% mean yield reduction across all cultivars. The greatest mean yield at 30.0 °C was observed in Bhrikuti (3279.0 kg ha⁻¹); a 25.5% reduction compared to the control at 10.0/25.0 °C. The lowest mean yield at 30.0 °C was observed in NL 1412 (754.5 kg ha⁻¹); the mean yield was reduced by 77.1% compared to the control (4144 kg ha⁻¹) at 10.0/25.0 °C.	^[11]
11 released varieties (India, Pakistan) 17 breeding lines (India, Pakistan) Chirya 3, BAZ (China)	Triticum aestivum L.	6.0/35.0	Post-anthesis.	59.5% mean yield reduction across all cultivars. The highest yielding (g/plant) genotypes during late sown heat stress were HD 2967 (5.7 g), BRW 3797 (5.4 g), and SW 129 (5.3 g) at 35.0 °C. The yield was reduced by 58.5%, 54.1%, and 47.7% for each cultivar, respectively. Raj 3765 displayed the greatest yield thermotolerance, reducing by 37.4% at temperatures up to 35.0 °C during post-anthesis.	^[32]
28 lines from International Maize and Wheat Improvement Centre (CIMMYT Mexico)	Triticum aestivum L.	8.0/35.0	Vegetative and grain filling.	50.0% reduction in yield from combined heat and drought stress. Borlaug 100 displayed stability, with a mean yield 8.0% higher than the control cultivars Roelfs F2007 and Vorobey.	^[31]
Yangmai16 (China) Xumai30 (China)	Winter Triticum aestivum L.	21.0/31.0 25.0/35.0 29.0/39.0 33.0/43.0	At anthesis and grain filling.	1.5–1.6% yield reduction per increase in heat degree days (above 30.0 °C) at anthesis, averaged across all treatments. 1.0–1.3% yield reduction per increase in heat degree days (above 30.0 °C) at grain filling, averaged across all treatments.	^[40]
Seri82 (CIMMYT Mexico)	Triticum aestivum L.	22.0/32.0	14 days at anthesis. 14 days during grain filling.	Grain yield per plant was reduced by 29.0% (from 3.5 g at the control temperature of 14.0/24.0 °C to 2.5 g at 22.0/32.0 °C) during anthesis. Grain yield per plant was reduced by 44.0% (from 3.5 g at the control temperature to 1.95 g at 22.0/32.0 °C) during grain filling.	^[14]
Almansor, Antequera, Bancal, Estero, Nabão, Pata Negra, and Roxo (Portugal)	Triticum aestivum L.	20.0/40.0	10 days after anthesis.	Bancal was the least affected variety, recording no significant difference in grains per the first spike between the control temperature (20.0/25.0 °C) and heat stress treatment (20.0/40.0 °C). Antequera was the most affected variety, recording a significant decrease in grains per the first spike, decreasing from 12 to 7 grains at the control and heat stress treatment, respectively.	^[41]

Similarly, Prasad and Djanaguiraman ^[42] identified the growth stages most sensitive to high temperature on the heat-sensitive cultivar Chinese Spring. Exposure to 5-d heat (26.0/36.0 °C) at 0–5 days pre-anthesis caused a maximum reduction in floret fertility (correlated to grain number) i.e., heat-stressed plants having ~25% fertility compared to the control (~80%) at 15.0/25.0 °C. Whilst heat stress during the period between 5–15 days pre-anthesis also had a significant impact on floret fertility, it had no significant impact on thousand kernel weight (TKW). In contrast, TKW was significantly decreased when 5-d heat (26/36.0 °C) was applied between 10 and 30 days post-anthesis; floret fertility was not impacted when heat was applied during this period, remaining at ~80%. In summary, heat stress at anthesis mainly lowers grain yield by reducing grain number per plant whereas heat stress post-anthesis (at grain filling) mainly lowers grain yield by reducing TKW and grain size.

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