Crop Physiology Case Histories for Major Crops

editors

Chapter 1: Maize

María E. Oteguia; Alfredo G. Cirilob; Sergio A. Uhartc; Fernando H. Andraded    a Department of Plant Production, School of Agriculture, CONICET at INTA Pergamino Experimental Station and University of Buenos Aires, Buenos Aires, Argentina

b Department of Crop Production and Environmental Management, INTA Experimental Station, Pergamino, Argentina

c Faculty of Agronomy, National University of the Northeast, Santiago del Estero, Argentina

d INTA-National University of Mar del Plata and CONICET, Balcarce, Buenos Aires, Argentina

Abstract

Maize is presently the grain crop that covers the largest cultivated area, has the largest production, and the largest grain yield worldwide. In recent years, maize occupied an important part of the land previously cropped to other species and moved forward into many newly cultivated areas considered fragile on a global scale. In all cases, it produced extraordinary economic benefits together with serious sustainability concerns. Considering this context and future climate scenarios, in this chapter, we revisit and upgrade the ecophysiological and agronomic concepts that helped maize conquer its current leading status among all cultivated species.

Keywords

Zea mays (L); Production systems; Grain yield; Resource use; Resource use efficiency; Biomass production; Kernel number; Kernel weight; Grain quality

1: Introduction

1.1: Global trends

Native to the New World, maize is the main staple for a large part of humankind, being especially important for several Latin American and African countries. In these countries, maize can be consumed as porridge (such as grits, polenta, or ugali), popcorn, roasted kernels, vegetable (fresh, frozen, or canned sweet maize), flour, or meal (bread, tortillas, chips, extruded snacks, etc.). Worldwide, its grains are used to produce ethanol (for beverages or as a fuel source), cooking oil, and starch. Grains and by-products are also used as animal feed, whereas its biomass is an energy source and is used as silage. Despite its broad distribution across all continents (Fig. 1.1a left), four of the top ten maize producing countries (Fig. 1.1b left) are in the Americas, where it covers the largest area worldwide. Maize is well-adapted around the world and returns high yields (Fig. 1.1c left). Low technological development sets a limit to crop yield in otherwise suitable environments (e.g. Africa).

Fig. 1.1 Left panels: world distribution (averaged for the 2008–17 period) of (a) area cropped to maize, (b) production, and (c) GY. Right panels: evolution of annual records of each variable for the 1961–2017 period. Lines represent (a and b) bilinear and (c) linear fitted models. The corresponding breakpoints (vertical arrows in a and b) and slopes (all variables) are indicated. Based on FAO, 2019. Records accessed on 1st Aug 2019, http://www.fao.org/faostat/en/#data/QC.

Land cropped to maize increased at a rate of close to 1 Mha y−  1 between 1961 and 2005 (Fig. 1.1a right) and increased to over 4 Mha y−  1 afterwards. A good part of the change has been driven by crop substitution, such as rice by maize in China. Variations in area produced similar variations in overall production (Fig. 1.1b right), with rates that increased from 10.5 to 34 M t y−  1 in almost the same period (breakpoint in 2003). Contrarily, global grain yield (GY) increased at a constant rate of 66 kg ha−  1 y−  1 for the whole period. Not surprisingly (Hall and Richards, 2013), outstanding breeding events such as the delivery of Bt corn to market in 1996 did not produce equivalent landmarks in maize production during the evaluated period.

Climate extremes, such as drought or heat stress, can lead to harvest failures and threaten the livelihood of agricultural producers and the food security of communities. Improving the understanding of their impacts on maize GY is crucial to enhance the resilience of the global food system. Climate factors, including mean climate and climate extremes, explain 16%–39% of the variance of yield anomalies (YA), with 10%–31% of the explained variance attributable to climate conditions (Fig. 1.2). YA related more closely with temperature extremes than with precipitation-related factors (Vogel et al., 2019).

Fig. 1.2 (a) Mean regional production shares per continent during 1990–2008 and standard deviation (SD) of production anomalies (after detrending) relative to mean production and (b) Explained variance of production shares accounted for yield anomalies (YA), mean climate conditions plus extreme events (MCC + EE), and mean climate conditions (MCC). Adapted from Vogel, E., Donat, M.G., Alexander, L.V, Meinshausen, M., Ray, D.K., Karoly, D., Meinshausen, N., Frieler, K., 2019. The effects of climate extremes on global agricultural yields. Environ. Res. Lett. 14, 054010, https://doi.org/10.1088/1748-9326/ab154b.

The forecast for future scenarios is a loss of climatic suitability for maize in Sub-Saharan Africa and Latin America regions but accompanied by an expansion in the northern hemisphere, particularly in Europe. The relative change in climatically suitable areas for future maize production was estimated for the top five producers. Production in 2050 is expected to increase 8% for the USA and 4% for China, and to decrease 5% for Brazil, 2% for Argentina, and 11% for México. The incidence of low temperature and waterlogging, presently common in Europe and Asia, is projected to diminish, whereas heat stress in Africa and drought stress in South America are projected to increase (Ramirez-Cabral et al., 2017).

1.2: Main production areas

The USA is the world leader in annual maize production, with 38.9 Mha (Fig. 1.1a) that produce 392 Mt (Fig. 1.1b). Twenty percent of the production is exported. After the introduction of teosinte in the USA territory thousands of years ago, many native communities adopted it as a staple crop as far north as Canada, reaching a production of ca. 13.9 M t y−  1. First European settlers to arrive to Western USA learned maize cultivation from native Americans and spread the crop eastwards. Presently, maize is produced in a variable proportion in most USA states, Iowa being the leader in total production and GY (Fischer and Edmeades, 2010), closely followed by Illinois and Nebraska.

China is the second global maize producer (Fig. 1.1b). Over the past few years, maize has become the main crop of the country, both in production (257 M t y−  1) and cultivated land (42.4 Mha−  1). This change is attributed to the increasing local demand for livestock feed (i.e. animal protein). A greater proportion of urban, wealthier population has driven consumption of meat products per capita to increase at a rate of 1.38 kg y−  1 since 2011. Over the past 25 years, maize production has increased 125%, while rice production has experienced a modest 7% increase. While in the 1940s, two-thirds of the Chinese maize crop production were used for human consumption, presently, 60% is used as animal feed, 10% for direct human consumption, and the remaining for industry.

Brazil is the third world maize producer, with nearly 94.5 M t annually and 17.3 Mha. This country is consolidating as the second main exporter. The production comes from two contrasting systems. One is called safra (full season), characterised by sowing starting early in August up to October–November. The other is called safrinha that presently occupies two-thirds of total area cropped to maize. The safrinha corresponds to late-sown maize (starting in January up to February), usually rotated with soybean to avoid fast soil nitrogen (N) decay. It predominates in the Mato Grosso states at the Centre-West of the country, which have displaced the traditional maize producing area located in the south–south east.

The fourth maize producer is Argentina, accounting for approximately 6.4 Mha and reaching 51 M t y−  1 in 2018–19. Argentina is presently the third exporter, close to Brazil. Like in the neighbour country, there are two main sowing dates: the early one that ranges between late August and early November and the late one from December to end of January. Delayed sowing became economically feasible with the introduction of Bt maize in 1997 (Otegui et al., 2002) and gained fast adoption after the severe drought of 2008–09. It presently occupies 45%–55% of the total area. Córdoba, in the geographical centre of the country, is presently the main maize producer province of Argentina.

The EU produces 61 M t of maize per year. France, chiefly in the south, leads maize production within the EU. The Aquitania region delivers 21% of overall France production, which is presently 14 M t y−  1 in 1.7 Mha. The crop is generally sown between April and May and is harvested between September and November. Owing to low local consumption, most of the French production is exported. Other important maize producers in the EU are Romania, Hungary, Spain, Germany, and Poland.

Ukraine, along with Russia, is the principal maize producer in the Black Sea region. Ukraine has vast sectors of black soils known as chernozems, which are among the most fertile in the world. Ukrainian growers produce 32 M t of maize annually in 4.3 Mha. In 2018–19, Ukraine accounted for 16% of global maize exports, becoming the fourth exporter.

India produces 28 M t of maize annually in 9.2 Mha. The crop is primarily grown in the northern states, including Uttar Pradesh, Bihar, Himachal Pradesh, and Rajasthan. Uttar Pradesh and Bihar account for nearly 16% and 14% of the country’s maize production, respectively. The crop is usually sown at the beginning of the rainy season, between mid-May and July, and is harvested between November and January.

Maize is the most important crop grown in Mexico, with almost 60% of the country’s cropland from sea level up to 2600 masl. Occident, Bajio, and Sinaloa regions contribute with nearly 60% of total production. Climate allows for two harvests per year, which in 2018–19 produced 27 M t in an area of 7.4 Mha. The first sowing takes place in spring–summer (April–July) and accounts for 75% of the annual production. The remaining 25% corresponds to the second one, which takes place during autumn–winter and is conducted under irrigation. Mexico has no export surplus but presently meets the internal demand of white maize for human consumption plus a small part for animal feed. However, it still needs to import 14 M t of maize for its growing livestock sector.

Indonesia is the leading maize producer (26 M t in 5.3 Mha) in the ASEAN Economic Community, followed by the Philippines (7.5 M t) and Vietnam (5.2 M t). Indonesia needs to import maize, primarily for livestock. The main constraint to Indonesian production is the lack of enough agricultural land, which is balanced with forested lands.

South Africa produced 16 M t of maize in 2.6 Mha during 2018–19. It is one of the main African producers together with Nigeria, Egypt, and Ethiopia (10 M, 8 M, and 7.8 M t, respectively). The crop is cultivated primarily in the north and north-east of the country. The South African provinces of Guateng, north-west, Mpumalanga, and Orange Free generate the highest maize GYs of the country. This crop is sown between September and December and is harvested between April and June.

The rest of the world (141 countries) accounts for the production of the remaining 140 M t to complete 1147 M t in 2018 (based on FAO, 2019).

1.3: Maize in rotations: Suitability of previous and consequences for following crops

Being a C4 plant, a high proportion of maize in the cropping sequence assures large biomass productivity and high water and radiation use efficiencies to the crop system (Caviglia et al., 2013). Maize has positive effects on soil carbon (C) balance and soil physical properties (e.g. infiltration, stability of aggregates) compared to other crops because it produces a high amount of stubble with a high C/N ratio. The effect of maize on soil carbon content and on the soil carbon content equilibrium depends on management practices and GY as well as on soil and climate. The annual soil carbon balance is positively related to maize GY and negatively associated with soil carbon content. For maize GYs higher than 10 t ha−  1 in the US Corn Belt, soil carbon balance was negatively associated with soil carbon percent, and the balance was positive at soil carbon percent below 3% (Lucas et al., 1977). For the south-east of the Pampas region of Argentina, crop sequences with maize caused smaller reductions in soil organic matter along the years when compared with rotations without maize (Studdert and Echeverría, 2002).

The most relevant short-term effects of the preceding crop in the rotation are related to biotic adversities and the availability of water and nutrients. Maize reduces N availability for the next crop because of its high nutrient demand and high total N harvested with the grain, which leaves a large amount of residues with a high C/N ratio. Accordingly, GY of non-fertilised wheat following maize is penalised and has a high response to N fertilisation when compared with wheat following other crops (Echeverría et al., 1992). Hence, a nitrogen (N) fixing crop or a crop with low N requirement is convenient after maize (Domínguez et al., 2001). Additionally, where water recharge of the soil profile during fallow and after sowing is limited and rainfall during the growing season is insufficient, the effect of the preceding crop on water availability becomes relevant. In this sense, maize usually consumes more water than the other summer crops (Della Maggiora et al., 2002).

Growing maize in consecutive seasons reduces GY (Berzsenyi et al., 2000). Repeating maize in the cropping sequence reduces its productivity when compared with maize in rotation after pasture or soybean (Domínguez et al., 2001; Farmaha et al., 2016; Sindelar et al., 2015). This positive rotation effect is usually because of the increased N supply to maize following soybean or pasture (Domínguez et al., 2001). Fertiliser, however, does not fully compensate the described positive crop rotation effect, which could be also attributed to (1) improvements in soil water availability, soil structure, and soil microbial activity, (2) reductions in weed, insect, and disease incidence, and (3) absence of phytotoxic compounds and/or presence of growth-promoting substances originated from crop residues (Bullock, 1992; Karlen et al., 1994). Nevertheless, in semi-arid regions like those in the north of Argentina, the rotation soybean-maize requires a minimum of 50% of maize to adequately supply carbon to the soil through residues and roots, improving water capture efficiency, soil physical characteristics, and tolerance to erosion (Martini and Angeli, 2017).

1.4: Multiple cropping

An option to intensify the use of agricultural land consists of sowing two or more crops per season as double crops, relay crops, or intercrops (Caviglia et al., 2004; Coll et al., 2012; Neto et al., 2010). These intensified systems have been implemented in many regions worldwide (Fischer et al., 2014). They increase annual land productivity and water, radiation, and nutrient productivity, mainly through improved resource capture (Andrade et al., 2015; Caviglia and Andrade, 2010). The increase in resource capture results in (1) reduced surface run-off, percolation, erosion, water contamination, and water table level and (2) increased stubble inputs, soil carbon content, soil protection, and better weed control. Including maize in intensified systems also ensures high resource use efficiency (Caviglia and Andrade, 2010) and biomass production (Monzon et al., 2014), which contribute to maintain soil carbon balance (Oelbermann and Echarte, 2011). Examples of intensified systems that include maize are (1) winter wheat–summer maize double cropping in the North China Plain, which provides about 33% of the nation’s maize production (Wang et al., 2012; Yu et al., 2006), (2) maize as a second crop after soybean harvest (safrinha) in Mato Grosso and Parana, Brazil, with more than 3.3 Mha in 2013, (3) maize grown as double crop after field pea or other winter crop as an alternative to the widespread wheat/soybean double crop in Argentina (Andrade et al., 2017; Mercau and Otegui, 2015), and (4) soybean as double crop after maize harvest or as relay intercrop from 40 to 60 days after maize sowing (Monzon et al., 2014).

Xu et al. (2020) estimated the worldwide average land equivalent ratio of the maize/soybean intercropping in 1.32 ± 0.02, underscoring the high potential of intercropping over sole crops. The productivity of such intensified option depends on the frost-free period, temperature regime, and water budget of each environment (Andrade and Satorre, 2015; Caviglia et al., 2013, 2019; Monzon et al., 2014; Van Opstal et al., 2011). The length of the growing season may limit the adoption of these alternatives in several regions. Accordingly, the increase in annual land productivity for maize–soybean double crop and relay intercrop was directly associated with the length of the growing season (Monzon et al., 2014).

2: Crop structure, morphology, and development

2.1: Main phenological events

Ritchie et al. (1986) summarised visual maize development in the most widely used phenological scale (Fig. 1.3a). The VT-R1 period (VT: tasselling; R1: silking) splits the crop between vegetative (Vn, with n representative of leaf number) and reproductive (Rn, with n spanning from visible silks in R1 up to physiological maturity in R6) phases. Shoot meristems, however, follow a microscopic differentiation pattern that was described and illustrated in detail for the first time by Bonnett (1940, 1948). This pattern includes (1) floral initiation (FI) of the main stem apex (Fig. 1.3a), which represents the shift from leaf primordia differentiation to male differentiation (FIM) and (2) FI at axillary meristems (Fig. 1.3a), which represents the shift from husks differentiation to female differentiation (FIF). The former will produce the panicle (described as tassel) responsible for pollen production. The latter will produce the spikes (described as ears), which will bear the ovaries (Fig. 1.3b). The FIM sets total leaf number (TLN) in the main stem and usually takes place when the number of visible leaf tips is one-third of TLN, reflecting the tight coordination between the number of leaves initiated by the apical meristem and the number of visible leaves up to FIM (Padilla and Otegui, 2005). This coordination holds across maize hybrids of different maturity groups, with TLNs between 15 (short maturity) and 25 (long maturity). There are no axillary buds in the topmost leaves between the apical meristem and apical ear bud because apical dominance arrests the acropetal differentiation of axillary buds at the time of FIM (Lejeune and Bernier, 1996). Therefore, FIF is delayed with respect to FIM (Otegui and Melón, 1997), and it occurs first at the last differentiated, topmost axillary bud in the so-called ear-leaf. The ear leaf is among the largest leaves of the plant and is located at approximately one-third of the TLN counting downwards along the stem (Dwyer et al., 1992). This proportion, however, is under genetic control and consequently, may vary across genotypes (Li et al., 2015), with the concomitant effect on leaf area distribution above and below the ear. The FIF continues downwards, which explains the existence of more than one grain-bearing ear per plant (i.e. prolificacy  >  1) depending upon the genotype (i.e. prolific vs. non-prolific), the resource availability per plant, and their interaction (Otegui, 1995; Tetio-Kagho and Gardner, 1988a). Buds at the lowermost nodes do not elongate and do not become reproductive. These buds may develop into tillers, particularly for some hybrids grown at very low stand densities in arid environments (Kapanigowda et al., 2010).

Fig. 1.3 (a) Schematic representation of the maize growth cycle. The x axis includes the scale proposed by Ritchie et al. (1986) together with cycle duration in thermal time (TT, °Cd) and in days after sowing (DAS) and main developmental events and phases. The span of the horizontal arrows represents (1) the period of crop response to temperature and photoperiod, and (2) the extension of the juvenile phase, the inducible phase, and the critical period for kernel number per plant (KNP) determination. Vertical arrows indicate the time of TLN and maximum leaf area index (LAIMAX) establishment, as well as of KNP, individual kernel weight (KW), and grain yield (GY) establishment. Data of cycle duration, leaf area index (LAI; m² of green leaves per m² of soil), and KW are representative of temperate hybrids with a TLN of ≈  20 and a relative maturity (RM) of ≈  120. Images of just extruded silks and a mature kernel with the characteristic black layer (BL) at the placental region are representative of the silking and physiological maturity stages, respectively. The latter is preceded by a schematic representation of a kernel at the half-milk stage (90%–95% of final KW), with the milk line (ML) separating the starchy from the milky endosperm. FIM: male floral initiation; FIF: female floral initiation. (b) Illustrative images of ear development between the start of active ear elongation (≈  V14) and silking (R1). In the example, the ear length increases from ≈  1.5 cm in (1) to ≈  8.5 cm in (6), depending upon the genotype and growing conditions. At the start of active ear growth, spikelets at the base of the ear have completely developed florets with silks of less than 1 mm (circle in 1). These silks will reach 14.5 cm at silking (the bottommost red arrow in 6). Floret development and silk elongation continues acropetally along the ear from (1) to (6). Active silk elongation can be observed in florets at the middle of the ear during 5–7 days before silking (4), when silks at bottommost positions are 4 cm long and have reached the tip of the ear. The bottommost ovaries will be 10 cm long during 1–2 days before silking (5), whereas those from the middle of the ear will reach silking on the same date with silks of only ≈  10 cm length (6). Almost all florets along the ear will reach complete floret development at silking. Pictures obtained by M.E. Otegui on maize crops grown a Pergamino (33° 56′ S, 60° 33′ W), Argentina. Reference scale is in mm.

From FI onwards, reproductive buds undergo the developmental processes that shape the characteristic morphology of maize tassel and ears (Bonnett, 1966). The final results of these processes are the events of anthesis (i.e. anthers dehiscence and pollen shed by tassels) and silking (silks extrusion from the husk by one or more ears per plant). At this stage, all differentiated leaves have completed their expansion and the crops have reached its maximum leaf area (Fig. 1.3a), whereas most florets along the ear of maize are totally developed and have started silk elongation (Fig. 1.3b). Contrary to other cereal crops as wheat, floret development per se does not represent a constraint to maize kernel set in most field conditions (Otegui et al., 1995a, 1995b; Otegui, 1997; Otegui and Melón, 1997; Rattalino Edreira et al., 2011), but environmental constraints that cause a delay in ear morphogenesis may affect the silking pattern (Rossini et al., 2012). A slight protandry (i.e. anthesis anticipation respect to silking of the topmost ear) is common in maize, producing an anthesis-silking interval (ASI) of a few days in standard growing conditions. This effect of apical dominance is enhanced when crops are exposed to stressful environments around flowering, increasing the ASI (Bolaños and Edmeades, 1993; D’Andrea et al., 2009; Edmeades et al., 1993; Hall et al., 1982), and reducing the total number of exposed silks (Debruin et al., 2018; Messina et al., 2019; Rossini et al., 2020) with the concomitant negative effect on kernel set. In contrast, some genotypes may express protogyny, i.e. anticipation of silking with respect to anthesis (Haskell, 1953), particularly when grown under no abiotic/biotic constraint at low stand density (Borrás et al., 2009). This condition usually improves pollination synchrony within and between ears (Uribelarrea et al., 2002), promoting increased prolificacy and kernel set (Cárcova et al., 2000). There is an important variation for prolificacy expression in maize (De Leon and Coors, 2002), with no clear benefit between prolific and non-prolific hybrids in many cropping conditions (Otegui, 1995). The exception to this trend is in drought-prone environments for which reduced stand density is recommended, where prolific hybrids have higher GY stability across years (Ross et al., 2020).

Ovary fertilisation along the ear proceeds between the start of silking (R1) and the start of active kernel growth of the early-fertilised ovaries (R2), which usually represents a limit to successful kernel set among the late-fertilised ones (Cárcova and Otegui, 2007). Early kernel growth is chiefly linked to water influx (Borrás et al., 2003b), whereas active biomass accumulation starts at R2 and continues up to R6 (Fig. 1.3a). Biomass accumulates steadily in kernels along this period in parallel with a steady decline in kernel water content. Maximum kernel weight (KW) is reached at physiological maturity (Fig. 1.3a), when a black layer of necrotic tissue is evident at the placental region of the kernel (Daynard and Duncan, 1969) and its water content has dropped to ≈  30%–40% (Borrás et al., 2004; Gambín et al., 2007). A good surrogate to anticipate R6 is to follow the ML development (Fig. 1.3a). This line represents the interface between the solid (starchy) and liquid (milky) matrices of the maturing endosperm (Afuakwa and Crookston, 1984) and progresses from the crown to the base of the kernel. A 50% ML (or half-milk stage) corresponds to ≈ 40% kernel moisture and ≈ 90%–95% of final GY, whereas there is no milk remaining when the black layer becomes evident. Anticipation of R6 is particularly important in the production of maize silage (Ma et al., 2006).

2.2: Genotypic and environmental drivers of maize development

2.2.1: Temperature

For a given genotype, the duration in days of the above-described cycle is primarily modified by temperature (Fig. 1.3a), which affects the extension of all developmental stages in most evaluated environments (Capristo et al., 2007; Kumudini et al., 2014; Shim et al., 2017; Tsimba et al., 2013). In the absence of photoperiodic effects and abiotic constraints, such as water or N deficiencies, maize developmental rate during the vegetative period (i.e. pre-VT) increases between a base temperature (Tb) of ≈  8–10 °C and an optimum temperature (Top) of ≈  30–35 °C, whereas temperatures >  Top reduce the developmental rate (Cicchino et al., 2010a; Gilmore and Rogers, 1958; Kiniry, 1991). References for Tb and Top are less accurate for the grain-filling period (Kumudini et al., 2014), being 8 °C in CERES-Maize model (Kiniry, 1991) but assumed as 0 °C by other authors (Sinclair et al., 1990; Tsimba et al., 2013). Within Tb and Top, the TT (in °Cd) requirement for completing a given phase is represented by the inverse of the slope in the model fitted to the response of the developmental rate to temperature; i.e. it is a genotype-dependent constant (Ellis et al., 1992). On the basis of the TT concept, several models have been developed to predict cycle duration that are used in crop simulation models such as CERES-Maize (Jones and Kiniry, 1986) and APSIM (Keating et al., 2003). These models differ in cardinal temperatures and the associated functions for computations. Therefore TT requirements reported in the literature must be taken with care for comparisons because they may vary depending upon the approach (Dwyer et al., 1999a; Kumudini et al., 2014). Predictions of these models usually focus on TT to anthesis, silking, and physiological maturity and do not include the period of kernel desiccation up to commercial harvest maturity. The only methods that include differences in kernel desiccation rate are the ones developed by FAO and the comparative RM or simply RM. Originally (Jugenheimer, 1958), the former used a kernel moisture content of 34%–36% as reference, which dropped to less than 20% in the 1990s (Marton et al., 2008). The latter is an adaptation of the originally known as Minnesota Relative Maturity Rating (MRMR) because of its origin (Peterson and Hicks, 1973), and it is the most widely used all across the Americas. The RM method ranks hybrids comparatively so that two RM units represent a 1% difference in kernel moisture at harvest. As a general reference, hybrids that range between 1014 and 1453°Cd for the whole cycle and represent RMs of 75–110 or FAO of 100–500 are recommended for latitudes between 48º and 39ºN, respectively, in Canada and the USA (Dwyer et al., 1999a; Troyer, 2001). The TT requirement rises to 1200–1700°Cd for the whole cycle (Fig. 1.3a) among hybrids recommended for mid-latitude environments (30–38ºS) as those representative of the Pampa region in Argentina (Capristo et al., 2007; Otegui et al., 1996), which correspond to RMs of 110–128 or FAO 500–600 (Di Matteo et al., 2016; Luque et al., 2006; Troyer, 2001). TT requirements (>  1500°Cd) and RMs (125–140) and FAO rankings (600–800) increase for subtropical and tropical conditions as those representative of Brazil (Bergamaschi et al., 2013; Tojo Soler et al., 2005) or India (Thimme Gowda et al., 2013). Despite this conceptual framework, TT requirements reported in literature must be taken with care for comparisons because they may vary widely depending upon the approach used for computation (Dwyer et al., 1999a, 1999b; Kumudini et al., 2014).

Expansion of all differentiated leaves takes place between crop emergence (VE) and tasselling-anthesis (Fig. 1.3a) at a relatively constant phyllochron of usually 37–42°Cd leaf−  1 among reported hybrids (Hesketh and Warrington, 1989) but may extend up to 75°Cd leaf−  1 among hybrids in the CERES-Maize database of DSSAT (Hoogenboom et al., 2017) and between 33 and 62°Cd leaf−  1 among inbreds (Giauffret et al., 1995). The TT requirement for leaf primordia differentiation at the apical meristem (i.e. plastochron) may range between 24.3 and 36.4°Cd leaf−  1, with Tb between 4.0 and 8.1 °C, depending upon the genotype and based on soil temperature at 5-cm depth (Padilla and Otegui, 2005).

2.2.2: Photoperiod

As a short-day plant (i.e. highest developmental rate under short photoperiod), the TLN may vary depending upon (1) the photoperiod during the inducible phase and (2) the degree of photoperiod sensitivity. Two sub-phases can be recognised during the vegetative phase of maize apical meristem, during which only leaf primordia are differentiated. The first one is a juvenile (i.e. photoperiod insensitive) phase, which is followed by an inducible (i.e. photoperiod sensitive) phase (Fig. 1.3a). The latter is subsequently followed by a realisation phase, when the meristem has already initiated reproductive differentiation and is no longer sensitive to photoperiod (Kiniry et al., 1983b). The FIM sets the limit between the inducible and realisation phases, and it is usually evident as an elongation of the apical meristem (Bonnett, 1966; Stevens et al., 1986). No similar morphological change identifies the transition between the juvenile and the sensitive phases of the meristem. Detection of a juvenile period in three photoperiod-sensitive hybrids was possible by exposing plants to alternating short and long photoperiods at different intervals during the cycle and recording tassel emergence date (Kiniry et al., 1983b). These experiments in controlled conditions allowed the detection of a juvenile phase that ended 4–8 days before FIM when plants were grown at short photoperiods (i.e. highly inducible). Therefore the inducible phase lasts 4–8 days under optimum, short photoperiods and is expected to be ≥  4–8 days for photoperiod-sensitive genotypes grown at photoperiods longer than a threshold. Subsequent experiments (Kiniry et al., 1983a) in controlled conditions allowed the estimation of the main parameters (Major, 1980) of maize developmental response to photoperiod (BVP: basic vegetative phase, sum of the juvenile and inducible phases under optimum photoperiods; TP: threshold photoperiod; PS: photoperiod sensitivity) for hybrids of different maturity groups. These parameters ranged between 139 and 344°Cd for BVP, 10 and 13.5 h for TP, and 0 and 36°Cd h−  1 for PS. The BVP (equivalent to earliness per se presently used in wheat, see Chapter 3: Wheat, Section 2.2) was the shortest for the early-maturity group (≤  139°Cd), and longest for the late-maturity group (≥  251°Cd). Duration of the juvenile phase presently documented at the CERES-Maize database of DSSAT range between 110 and 458°Cd, whereas photoperiod sensitivity ranges between 0 and 5 d h−  1 (Hoogenboom et al., 2017). The use of days for the inducible phase allows for a variable TT duration of this phase, which may or may not modify TLN depending upon temperature. This subtle change is key to capture genotype × environment effects that are usually evident as a variation in the TLN in field conditions because of variable temperature regimes (Tollenaar and Hunter, 1983). Such effects are commonly registered even for the same hybrid sown in the same site and sowing date across years (Cirilo and Andrade, 1994a; Otegui et al., 1995b).

Knowledge of the previously described genotype × temperature × photoperiod interactions is critical for matching genotype and environment under two main premises. Firstly, total cycle duration must fit within the length of the growing season and must avoid excessively low temperatures at early and late growth stages. These situations are frequent in exceptionally early (Otegui et al., 1996) and late sowings (Bonelli et al., 2016; Mercau and Otegui, 2015). In extremis, early frosts may cause seedling mortality and reduce stand density and uniformity because maize cannot compensate for plant losses. Uneven stands are particularly critical in maize (Liu et al., 2004) because they promote hierarchies among plants (i.e. dominant and dominated individuals) with negative effects on the final number of kernels m−  2 and GY (Maddonni and Otegui, 2004). Late frost may prematurely arrest grain filling with direct penalisation of GY (Baum et al., 2019; Bonelli et al., 2016; Mercau and Otegui, 2015). Such penalisation may also take place with excessively low irradiance late in the cycle, which reduces plant growth per set kernel (Cirilo and Andrade, 1996; Maddonni et al., 1998), may hasten reserve deployment (Jones and Simmons, 1983; Uhart and Andrade, 1991) and may shorten the grain-filling period and increase grain moisture at physiological maturity (Sala et al., 2007). Secondly, sowing date and genotype selection should pursuit the concurrence of the critical period for kernel set with the optimum conditions for crop growth (Section 4.1) in order to (1) avoid negative effects of the environment on the ASI (in days), which affects kernel set (Bolaños and Edmeades, 1993; Hall et al., 1982), and (2) maximise the duration of the critical period for kernel set (Otegui and Bonhomme, 1998) and crop growth rate (CGR) during the critical period (Andrade et al., 1999) for maximising the number of kernels m−  2 (Section 4.1).

3: Growth and resources

3.1: Capture and efficiency in the use of radiation

3.1.1: Canopy size and light interception

The total shoot biomass (BT) produced in a period of n days can be expressed as the cumulative daily product of incident photosynthetically active radiation (IPAR), the efficiency of the canopy for intercepting IPAR (ei), and the radiation use efficiency (RUE) (Eq. 1.1).

   (1.1)

During early growth, maize canopies invest a large proportion of resources (photoassimilates and nutrients) in leaves, which in turn promotes ei. From seedling emergence (VE) to flowering (VT-R1), the canopy will develop a leaf area three to five times greater than the land area it covers, a relationship known as LAI (Fig. 1.3a). In the absence of plant stress, CGR increases with increasing ei, peaking at ei = 0.95. The LAI required to achieve this ei is the critical LAI (LAIC). In maize, a typical LAIC is 3–4. Differences in LAIC can be related to contrasting leaf habit (e.g. erectophile or planophile) between hybrids (Maddonni and Otegui, 1996) and sowing patterns (Maddonni et al., 2001, 2006).

Then, ei is an exponential function of LAI, with a curvature that depends upon an extinction coefficient k (Eq. 1.2) that usually ranges between 0.4 and 0.6 for maize.

   (1.2)

Leaf area is a function of leaf area growth and senescence. Leaf area growth depends on leaf number and size. Leaf number is mainly regulated by the genotype (Dwyer et al., 1992) and by the environmental conditions, depending on the crop response to photoperiod and temperature (Section 2.2). Leaf area per plant may be greater under long photoperiods that favour more leaves in sensitive hybrids (Bonhomme et al., 1991; Kiniry et al., 1983b). Individual leaf development and expansion, however, are mainly driven by temperature (Section 2.2; Ritchie and Nesmith, 1991).

Leaf appearance rate increases between 8 and 34 °C and decreases at higher temperature (Kiniry, 1991). Leaf expansion rate (LER) is a direct function of temperature (greatest between 22 and 32 °C; Ritchie and Nesmith, 1991), whereas duration of the expansion period is inversely related to temperature (Hay and Walker, 1989). Owing to total or partial compensation between these two processes, the canopy achieves maximum leaf area when the average daily temperature is 21 °C (Hardacre and Turnbull, 1986; Reid et al., 1990; Wilson et al., 1973). A consequence of these characteristics is the increase in LAI in response to the delay in sowing date registered for spring sowings among temperate hybrids grown at the same stand density under no abiotic constraint (Maddonni and Otegui, 1996). LER is highly sensitive to water and nutrient deficits, as discussed in Sections 3.2.3 and 3.3.2. From crop emergence to anthesis, generation and expansion of leaves dominate leaf area dynamics, with little contribution of senescence except in extreme situations (e.g. high plant population density and shortage of N). Early vigour and maintenance of active leaf area ensure high capture of radiation and thus high biomass production. Maximum green leaf area is attained at anthesis (Fig. 1.3a). Afterwards, changes in leaf area are a direct function of the senescence rate (Borrás et al., 2003a).

Leaf senescence is the series of biochemical and physiological events comprising the final stage of development, from the fully expanded state until leaf death. Leaf senescence is genetically determined and modulated by environmental factors (Thomas and Ougham, 2015). For instance, maize expressing the sunflower transcription factor HaHB11 exhibits improved GY chiefly through delayed leaf senescence (Raineri et al., 2019). Key environmental factors accelerating senescence include low radiation, low red:far-red ratio, water and nutrient deficiencies, vascular and leaf diseases, and unbalanced source–sink ratio (Borrás et al., 2003a; Rajcan and Tollenaar, 1999; Sadras et al., 2000).

3.1.2: Radiation-use efficiency and its response to environmental factors

Maize average RUE during the season is higher than for other summer grain crops such as soybean and sunflower (Andrade et al., 2005). For maize growing under adequate conditions, RUE is between 2 and 4 g MJ−  1 (Andrade et al., 1992; Hao et al., 2016; Lindquist et al., 2005; Otegui et al., 1995b; Westgate et al., 1997), when obtained from spot measurements at midday on sunny days and expressed on shoot biomass and a photosynthetically active radiation (PAR) basis. The superior RUE of maize results from (1) its C4 metabolism (Hesketh, 1963), with leaf photosynthetic rate 30%–40% higher than C3 species such as soybean, (2) its lower extinction coefficient (Eq. 1.2) that allows a more uniform distribution of the incoming radiation within the crop canopy, and (3) the low energy cost of carbohydrate-rich plant tissues (cellulose during vegetative growth and starch during grain filling) compared to protein and fat-rich tissues (Varlet-Grancher et al., 1982).

Sinclair and Muchow (1999) and Stöckle and Kemanian (2009) reviewed the sources of variation in RUE for different species. Low temperatures (Andrade et al., 1993; Westgate et al., 1997), water deficits (Muchow, 1989a), and nutrient deficiencies (Muchow and Davis, 1988; Uhart and Andrade, 1995a) reduce RUE. In most crop environments, low temperatures during early vegetative growth in early sowing and during grain filling in late sowing reduce RUE (Cirilo et al., 1992; Cirilo and Andrade, 1994a; M.E. Otegui et al., 1995b; Wilson et al., 1995)(Cirilo et al., 1992; Cirilo and Andrade, 1994a; Wilson et al., 1995). The effect of water and nutrient deficiencies on crop biomass production, however, is largely accounted for by the reduction in the amount of radiation intercepted by the crop because the decline in RUE is generally less important (Boyer, 1970; Gifford et al., 1984; Uhart and Andrade, 1995a). This is the consequence of leaf expansion being much more sensitive to water and nutrient deficits than photosynthetic rate per unit leaf area (Muller et al., 2011; Sadras and Milroy, 1996; Salah and Tardieu, 1997). Reductions in RUE as a result of water deficits are explained by stomatal or non-stomatal factors (Farquhar and Sharkey, 1982), according to the direction of the changes in CO2 concentration in the stomatal cavity (decreases or increases, respectively). High source–sink ratio during grain filling can reduce RUE (Borrás and Otegui, 2001; Rajcan and Tollenaar, 1999), possibly because of photosynthetic feedback inhibition.

3.1.3: Crop growth rate and growth duration in response to management practices

As it was already indicated, CGR is a function of IPARi and RUE, which depend on temperature and on the water and nutrient status of the crop. In turn, growth duration is determined by factors controlling phenological development, mainly temperature (Section 2.2). Rate of growth and growth duration are integrated into conceptual variables largely correlated with total biomass accumulation, i.e. photothermal quotient and growth per unit TT (Andrade et al., 1999; Fischer, 1985).

Maize canopies, as in all annual species, do not profit from all the IPAR during the growing season. The proportion intercepted generally ranges between 59% and 79% of total IPAR (Otegui et al., 1995b). This limitation can be overcome in part by (1) early sowing, which does not improve the proportion of IPAR that is intercepted along the season but improves the total amount of light intercepted by the crop (Bonelli et al., 2016; Cirilo and Andrade, 1994a; Otegui et al., 1995b); (2) long season hybrids (Otegui et al., 1995b); (3) increasing plant population density (Westgate et al., 1997), and (4) reducing row spacing (Andrade et al., 2002a; Maddonni et al., 2006). These practices increase IPARi because they promote rapid canopy closure and/or increase the amount of IPAR. The benefits of early ground cover do not translate into increased GY if they do not improve ei at the critical stages of GY determination (Maddonni et al., 2006; Westgate et al., 1997; Section 4.1).

Delaying sowing hastens vegetative growth and development because of high temperatures. Vegetative growth, however, is accelerated to a greater extent, so late-sown plants are generally larger than those sown early (Cirilo and Andrade, 1994a; Knapp and Reid, 1981; Maddonni and Otegui, 1996). Under these conditions, crops achieve maximal light interception in a shorter period from emergence (Bonhomme et al., 1994; Cirilo and Andrade, 1994a; Maddonni and Otegui, 1996). However, shortening of the growing cycle in late sowings decreases the total amount of radiation intercepted by the crop and, thus crop dry matter at harvest (Cirilo and Andrade, 1994a; Otegui et al., 1995b; Srivastava et al., 2018). Delays in sowing result in deterioration of some environmental conditions (i.e. reduced incident radiation and reduced RUE related to reduced temperatures) during the critical period for grain number determination and mostly during grain filling (Bonelli et al., 2016; Cirilo and Andrade, 1994b; Tsimba et al., 2013; van Roekel and Coulter, 2011), as analysed in Section 4.

Plant density is the practice with the greatest impact on LAI and hence on light interception of maize canopies (Overman and Scholtz, 2011). LAI decreases markedly in response to reductions in plant density (Cox, 1996; Maddonni et al., 2001; Tetio-Kagho and Gardner, 1988b) because leaf area per plant does not vary much when resources per plant increase (Andrade et al., 2005). This lack of vegetative plasticity in maize is the consequence of a very stable leaf size, a nearly constant leaf number (Vega et al., 2000), and a low capacity for tillering (Doebley et al., 1997). Thus radiation interception in maize is highly responsive to plant density (van Roekel and Coulter, 2011). This decrease in ei with reduced plant densities contrasts with the response of other crops. Decreasing plant density results in large reductions in IPARi at the critical period for grain number determination in maize, which results in reduced CGR at flowering and, in turn, in lower number of grains per unit area (Andrade et al., 1999).

Decreasing row spacing at equal plant densities leads to more equidistant plant distribution, hence reducing plant-to-plant competition for water, nutrients, and light, and increases intercepted radiation and biomass (Barbieri et al., 2008, 2012, 2013; Bullock et al., 1988). It also reduces the LAI required to intercept 95% of the incident radiation because of a higher light extinction coefficient (Flénet et al., 1996; Riahinia and Dehdashti, 2008). In the absence of significant water and nutrient deficiencies, however, the benefits of decreasing row spacing are variable. Some researchers report GY increases (Bullock et al., 1988; Olson and Sander, 1988; Scarsbrook and Doss, 1973), while others do not (Ottman and Welch, 1989; Westgate et al., 1997; van Roekel and Coulter, 2012). GY responses to decreased distance between rows are inversely proportional to ei achieved with the wide row control treatment during the critical period for grain number determination (Andrade et al., 2002a). Thus GY increase in response to narrow rows is closely related to the improvement in ei during the critical period for grain set.

Full light interception can probably not be achieved when (1) short-season and/or erect-leaf cultivars are grown (Bavec and Bavec, 2002); (2) plants are defoliated by frost, hail or insects; or (3) plants are subjected to water or nutrient stress at vegetative stages (Barbieri et al., 2000). Because drought or nutrient deficiencies during vegetative periods limit leaf area expansion (D’Andrea et al., 2006; Salah and Tardieu, 1997; Uhart and Andrade, 1995a), they would increase the probability of response to reduced row spacing. Early sowing in maize could also increase the response to reductions in row spacing because this practice leads to smaller plants with fewer leaves (Duncan et al., 1973; Maddonni and Otegui, 1996) (Section 2.2).

The length of the growing cycle is critical in matching genotype and environment (Capristo et al., 2007; Wilkens et al., 2015). In general, the longer the growing season, the longer the maturity group of adapted cultivars (Section 2.2). At low latitudes, temperature and radiation do not vary much along the year, and long-season hybrids are generally more suitable because they compensate reductions in cycle duration because of high mean temperature with more leaves (Section 2.2); this phenotype enables to capture more incoming radiation than short-maturity hybrids in those environments (Lafitte and Edmeades, 1997). Contrarily, at high latitudes, radiation and temperature decrease markedly during grain filling (Maddonni et al., 1998). Therefore, GY usually decreases when sowing is delayed and hybrid maturity class is increased above the limit set to total cycle duration by the frost-free period (Baum et al., 2019). A short-season hybrid with low leaf area per plant and low vegetative plasticity may not achieve full light interception at the critical stages (Eq. 1.2) and therefore is more likely to benefit from higher plant density and reductions in row spacing than a long-season cultivar (Assefa et al., 2016; Lindsey and Thomison, 2016; Sarlangue et al., 2007).

The detrimental effects of delayed sowing in maize are, in general, more pronounced in long-season hybrids. These hybrids benefit most from early sowings and show the largest reductions in GY in response to delayed sowing (Olson and Sander, 1988; Tsimba et al., 2013). The benefit of late-sown early-maturity hybrids depend on the magnitude of the delay and the potential length of the growing season (Baum et al., 2019; Lauer et al., 1999).

3.2: Capture and efficiency in the use of water

3.2.1: Environmental patterns of water supply and demand

Within the broad range of temperatures and frost-free periods for maize outlined in Section 1, crops are mostly grown where rainfall exceeds the 250 mm y−  1, with no rainfed commercial production and with rainfall below 150 mm during the warm season (Shaw, 1988). This distribution is linked to the high sensitivity of maize GY to water deficits. For instance, Meng et al. (2016) estimated a decline of 0.17% mm−  1 in relative GY (i.e. quotient between rainfed and irrigated GY) when rainfall dropped below 462 mm in the Chinese maize belt during the growing season, with no GY record for rainfall ≤  240 mm. Globally, Rattalino Edreira et al. (2018) estimated a marked decline (64%) in rainfed water productivity (i.e. GY per unit of potential crop evapotranspiration under water-limited conditions) when average evaporative demand increased from 3 to 7 mm d−  1 or when the fraction of soil evaporation to potential rainfed crop evapotranspiration increased from less than 20% to more than 40%. Both low evaporative demand and low soil evaporation usually corresponded to humid, cool high-latitude environments typically represented by European countries and the north-central USA. In contrast, high values corresponded to arid, warm low-latitude environments well represented by sub-Saharan Africa and the western US corn belt.

It has been well documented that maize GY reductions are particularly pronounced when water deficits take place around flowering (Claassen and Shaw, 1970; Hall et al., 1982). In these conditions, GY can be more affected than biomass production, bringing a marked decline in harvest index (HI, grain to total biomass ratio) and water productivity (Sinclair et al., 1990). Despite these characteristics, Rattalino Edreira et al. (2018) estimated that maize global GY gaps because of non-water limitations (i.e. the difference between potential water-limited GY and on-farm GY) could be equivalent to those produced by water scarcity (i.e. the difference between potential GY and potential water-limited GY), even in climatic zones traditionally considered as highly water-limited as the sub-Saharan Africa. This paradigm shift is expected to modify, at least partially, systems analysis of crop management to improve water productivity.

3.2.2: Root expansion and senescence, root size, architecture, and functionality

Typical for gramineous plants, maize main root system corresponds to nodal or crown roots that arise from basal nodes (i.e. older) (Gregory, 2006). These nodes do not elongate and produce the adventitious root system that comprises multiple root axes and their laterals. The system is of fibrous, finely distributed appearance. Additional aerial roots from higher nodes may grow into the soil and contribute to water and nutrient uptake and plant anchorage. McCully (1999) summarised the characteristics of field-grown maize roots, indicating that first-order laterals are short (less than 3 cm) when compared with lab-grown roots, most of them reach final length in less than 2.5 days, and they usually persist along the whole cycle. In this system, however, maturation for adequate water transfer does not take place until 15–40 cm from the root tip for large xylem vessels, whereas for small vessels, this distance is reduced to 4–9 cm and for the very narrow protoxylem to about 1–2 cm. Tips of lateral roots have early and rapid senescence, which progresses towards its older, basal part (Fusseder, 1987). Although only one-third of these roots produce second-order laterals, overall laterals represent × 30 the length of the axial roots. In this system, root hairs are key to water and nutrient uptake, and their life-span ranges between a few days and 1–3 weeks (Fusseder, 1987), depending upon the method used for the analysis (cytoplasmic intactness or nuclear staining, respectively).

The profuse maize root system develops primarily between sowing and R2 (McCully, 1999), when it reaches its maximum depth (Fig. 1.4a). For soils with no permanent limitation to root growth, the evolution of root depth follows a general sigmoid pattern (Fig. 1.4a). Higher soil bulk density (e.g. silty clay loam soils of the Argentine Pampas respect to loam soils of Iowa) and reduced temperature (high latitude with respect to low latitude) may delay maize root penetration across soil layers, modifying the root front velocity (RFV; Fig. 1.4a). Estimated maximum RFVs for maize ranged between 2.4 and 3.4 cm d−  1 and were reached between 43 and 56 days after sowing in the evaluated temperate environments. In a given environment, hybrids with extended cycle duration (i.e. high RM) are expected to have deeper roots than their short-cycle counterparts, whereas those with acute root angles are expected to have enhanced RFV with respect to those with wide angles (Hammer et al., 2009). Maximum root depth (Fig. 1.4b) and root abundance at each soil layer (Fig. 1.4c) vary with growing conditions. For instance, maize rooting depth can reach 2.25 m on loam soils (Typic haplustols) when exposed to terminal drought in a mid-latitude environment (Dardanelli et al., 1997). In contrast, no water uptake was detected below 1.65 m under similar drought conditions when crops were growing on silty clay loams (Typic argiudols) of the same region (Fig. 1.4b). Roots can traverse dense layers when layers are wet or bypass them through cracks when soils dry (Dardanelli et al., 2004). However, root proliferation within a soil layer is severely affected by increased bulk density (Fig. 1.4c), which promotes root clumping and the concomitant negative effect on water extraction (Fig. 1.4b and d). This restriction may also compromise root proliferation in subsequent soil layers (Dardanelli et al., 2004), which may not reach the critical root density for maximum water extraction rate, reducing the actual amount of extractable soil water in deep soil layers (Carretero et al., 2014). Similarly, management practices with a negative effect on soil structure also affect maize root proliferation, as observed for long respect to short cropping periods in a rotation (Cárcova et al., 2000) and for soil densification related to tillage (Díaz-Zorita et al., 2002; Taboada and Alvarez, 2008).

Fig. 1.4 Maize root system: expansion, size, and functionality. (a) Evolution pattern of the root front of maize crops growing in different environments (lower panel) and derived RFV (upper panel). The former was estimated from depletion of soil water (Argentina) or in situ measurements (USA). DAS: days after sowing. (b) Maximum rooting depth based on relative plant available soil water (PASW) at physiological maturity ( dashed line ) of maize crops grown on contrasting soil types ( circles : Haplustol soil; squares : Typic argiudol soil) in the Pampas of Argentina. The solid lines correspond to initial soil water content (0: permanent wilting point; 1: field capacity). (c) Root abundance (as proportion of in situ evaluated soil blocks with at least one visible root) at R1 of one maize hybrid grown on different soils of central France. Vertical dashed lines indicate the presence of massive structures (arenosols and planosols) or shrinkage cracks (vertisols). (d) Proportion of total water uptake from different soil layers during the silking-20 days ( solid lines ) and silking  +  20 days ( dashed lines ) periods of maize crops exposed to contrasting water regimes. a: Adapted from Cárcova, J., Maddonni, G.A., Ghersa, C.M., 2000. Long-Term Cropping Effects on Maize on Maize: Crop Evapotranspiration and Grain Yield. Agron. J. 92, 1256–1265. https://doi.org/10.2134/agronj2000.9261256x; Otegui, M.E., 1992. Incidencia de una sequía alrededor de antesis en el cultivo de maíz. Consumo de agua, producción de materia seca y determinación del rendimiento. Tesis MSc. Facultad de Ciencias Agrarias, Universidad Nacional de Mar del Planta; Ordóñez, R.A., Castellano, M.J., Hatfield, J.L., Helmers, M.J., Licht, M.A., Liebman, M., Dietzel, R., Martinez-Feria, R., Iqbal, J., Puntel, L.A., Córdova, S.C., Togliatti, K., Wright, E.E., Archontoulis, S. V., 2018. Maize and soybean root front velocity and maximum depth in Iowa, USA. F. Crop. Res. 215, 122–131. https://doi.org/10.1016/j.fcr.2017.09.003; b: Adapted from Dardanelli, J.L., Bachmeier, O.A., Sereno, R., Gil, R., 1997. Rooting depth and soil water extraction patterns of different crops in a silty loam haplustoll. F. Crop. Res. 54, 29–38. https://doi.org/10.1016/S0378-4290(97)00017-8; Otegui, unpublished; c: Adapted from Nicoullaud, B., King, D., Tardieu, F., 1994. Vertical distribution of maize roots in relation to permanent soil characteristics. Plant Soil 159, 245–254. https://doi.org/10.1007/BF00009287; d: Adapted from Otegui, M.E., Andrade, F.H., Suero, E.E., 1995a. Growth, water use, and kernel abortion of maize subjected to drought at silking. F. Crop. Res. 40, 87–94. https://doi.org/10.1016/0378-4290(94)00093-R.

3.2.3: Crop water use and canopy conductance as related to canopy architecture, stomatal conductance, and canopy-atmosphere coupling

Common to all crops, biomass production (Eq. 1.1) and the amount of water transpired by maize canopies are tightly related to the amount of solar radiation (SR) intercepted along the cycle (McNaughton and Jarvis, 1991), and both processes depend upon leaf differentiation (Section 2) and expansion (Fig. 1.5). Crop capacity to intercept light (Eq. 1.2) depends chiefly on a high leaf elongation rate (LER) to achieve the critical LAI (LAIC; Section 3.1.1) at the start of the critical period for kernel set (Section 4.1; Fig. 1.5d and e). In these conditions, LER depends on temperature in the 10–35 °C range (Sadok et al., 2007) and varies with leaf position along the stem (Andrieu et al., 2006; Dwyer et al., 1992). Therefore and provided all other variables remain uniform (e.g. VPD, mineral nutrition, stand density), the effects of soil water deficits (Fig. 1.5a and c) on maize LER (Fig. 1.5c, inset; Tanguilig et al., 1987) and LAI (Fig. 1.5d; Cavero et al., 2000) are directly driven by fluctuations in the soil–leaf water potential continuum that affect cell turgor (Boyer, 1970; Reymond et al., 2003) and plant hydraulic conductivity. These fluctuations depend primarily upon the crop water regime (Fig. 1.5a) and can be influenced by management practices (row spacing, tillage system, soil mulching) that affect the partitioning of crop evapotranspiration (ETc) between transpiration (T) and soil evaporation (Es) (Al-kaisi et al., 1989; Villalobos and Fereres, 1990). Reductions in intercepted SR because of water shortages will depend upon the extent of effects driven by LAI reductions on ei (Fig. 1.5e), which include the pre-silking period of canopy expansion and canopy senescence (Fig. 1.5d).

Fig. 1.5 Maize crop water use. DSSAT 4.7.0.0 ( Hoogenboom et al., 2017) simulated results of processes conducive to the amount of water transpired by crops grown under contrasting water regimes in Spain (based on Cavero et al., 2000). DAS: days after sowing. (a) Water regimes and their effects on the fraction of transpirable soil water (FTSW), (b) crop (ETc) to potential (PET) evapotranspiration ratio, (c) degree of water stress (SI: stress index) on tissue expansion processes, and (d) LAI along the cycle. Effects on LAI summarise the response of LER to pre-dawn leaf water potential (Ψ) (c inset; Chenu et al., 2008). (e) Large effects on LAI did not affect to the same extent maximum light interception efficiency (ei; Eq. 1.1) of stressed crops but (f) did affect their capacity to hold it along the reproductive phases, with the concomitant variation in the amount of SR intercepted by canopies that drive plants transpiration. Short black arrows in (a–e) indicate silking date on ca. 80 DAS. Squares in (d) correspond to observed LAI at silking and maturity. Long vertical arrows in (d) and (e) indicate the date when ‘control’ and ‘water deficit 1’ plots reached ≈  95% ei, which corresponded to LAI ≅ 4.5 (horizontal arrow in d). Plots under water deficit 2 reached a maximum LAI of 3.45 and ei = 0.87.

Water deficit reducing LAI reduces both cumulative intercepted SR (Muchow, 1989a) and transpiration (Muchow and Sinclair, 1991; Tanguilig et al., 1987). The relative response, however, differs among water regimes (Fig. 1.5f), being larger for water loss (up to 57% in the example) than for SR captured (up to 32% in the example). This uneven response is because of the additional reduction of transpiration in response to the decline in canopy conductance (gc) in drying soils. As an isohydric species (Tardieu and Simonneau, 1998), maize tends to keep a high leaf water potential (Ψ) by an efficient stomata control through hydraulic and non-hydraulic signals. The latter is mediated by xylem abscisic acid (ABA) (Bahrun et al., 2002), for which there is still some controversy about its origin in the plant (Christmann et al., 2007; Jackson, 1997). Main effects of increased xylem ABA on growth of maize plants exposed to water deficits are related to stomatal control (Tardieu et al., 2010). Increased ABA reduces leaf expansion (Cramer and Quarrie, 2002) and enhances stomatal closure, both conducive to reduced transpiration. Stomatal closure reduces gc, which maintains the plant water status (i.e. prevents a large drop inΨ) and improves plant hydraulic conductivity. Moreover, ABA effects on overall improved water status seem partially mediated by enhanced aquaporins activity, as reported by Parent et al. (2009) for maize lines divergently transformed in the NCED (9-cis-epoxycarotenoid dioxygenase) gene. Plants of ‘sense lines’ (i.e. those with enhanced ABA production with respect to the ‘antisense’ ones) had improved expression of mRNA for aquaporin PIP genes and related protein contents.

Variations in gc may also relate to fluctuations in atmospheric vapour pressure deficit (VPD). Increased VPD can promote stomatal regulation through upregulation of foliar ABA biosynthesis triggered by reduced leaf turgor (McAdam and Brodribb, 2016). Maize hybrids vary in transpiration response to both soil water content (Fig. 1.6a) and VPD (Fig. 1.6b) (Gholipoor et al., 2013).

Fig. 1.6 Maize soil–plant–atmosphere coupling and water use efficiency. Transpiration response of two maize hybrids to the (a) fraction of transpirable soil water (FTSW) and (b) vapour pressure deficit (VPD). EOT , end of treatment. Adapted from (a) Ray and Sinclair (1997) and (b) Gholipoor et al. (2013). (c) Response