post anthesis meaning

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INTRODUCTION

This topic serves as an overview for post-anesthetic care and the most common problems encountered in the post-anesthesia care unit (PACU). Preoperative evaluation and preventive strategies are discussed elsewhere. (See "Preoperative medical evaluation of the healthy adult patient" and "Management of cardiac risk for noncardiac surgery" and "Evaluation of perioperative pulmonary risk" .)

PHASES IN THE POST-ANESTHESIA CARE UNIT

Initial handoff  —  The initial handoff from the anesthesia care team to personnel in the PACU is typically standardized, as shown in the examples ( figure 1 and table 1 ). Further discussion is available in separate topics. (See "Handoffs of surgical patients" and "Patient handoffs" .)

Phase I and II care  —  PACU care is typically divided into two phases. Phase I emphasizes ensuring the patient's full recovery from anesthesia and return of vital signs to near baseline. Phase II recovery focuses on preparing patients for hospital discharge, including education regarding the surgeon's postoperative instructions and any prescribed discharge medications.

First days of anthesis: C. texana

First days of anthesis: c. carduacea, last days of anthesis: c. texana, last days of anthesis: c. carduacea.

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[ an- thee -sis ]

• the period or act of expansion in flowers, especially the maturing of the stamens.

/ ænˈθiːsɪs /

• the time when a flower becomes sexually functional

/ ăn-thē ′ sĭs /

• The period during which a flower is fully open and functional.
• Also called efflorescence

Discover More

Word history and origins.

Origin of anthesis 1

Example Sentences

These swellings help to spread out the branches especially at the time of anthesis.

Relation of temperature to anthesis and blossom drop of the tomato together with a histological study of the pistils.

Anthesis, the period or the act of the expansion of a flower.

These glands secrete a viscid juice at the time of anthesis.

Anthesis, anthropocosmic—— Say, I'm glad you didn't call me that!

postanthesis

• 1.1 Alternative forms
• 1.2 Etymology
• 1.3.1 Antonyms

English [ edit ]

Alternative forms [ edit ].

• post-anthesis

Etymology [ edit ]

From post- +‎ anthesis .

Adjective [ edit ]

postanthesis ( not comparable )

• occurring after the opening of a flower

Antonyms [ edit ]

• preanthesis

• English terms prefixed with post-
• English lemmas
• English adjectives
• English uncomparable adjectives

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anthesis noun

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Earlier version

• anthesis in OED Second Edition (1989)

What does the noun anthesis mean?

There is one meaning in OED's entry for the noun anthesis . See ‘Meaning & use’ for definition, usage, and quotation evidence.

How common is the noun anthesis ?

How is the noun anthesis pronounced, british english, u.s. english, where does the noun anthesis come from.

Earliest known use

The earliest known use of the noun anthesis is in the late 1700s.

OED's earliest evidence for anthesis is from 1783, in C. Linnaeus' Syst. Veg.

anthesis is a borrowing from Latin.

Etymons: Latin anthesis .

Nearby entries

• antheridium, n. 1818–
• antheriferous, adj. 1799–
• antheriform, adj. 1802–
• antherine, n. 1689–
• antherless, adj. 1798–
• antherogenous, adj. 1847
• antheroid, adj. 1818–
• antherozoid, n. 1853–
• antherozoidal, adj. 1865–
• anther valve, n. 1839–
• anthesis, n. 1783–
• anthias, n. 1601–
• anthill, n. Old English–
• ant-hillock, n. 1656–
• ant-hilly, adj. 1796–
• anthine, n. & adj. 1601–1768
• ant-hive, n. 1817–
• antho-, comb. form
• anthobian, n. & adj. 1835–
• anthocarpous, adj. 1835–
• anthocephalous, adj. 1847

Meaning & use

The Anthesis [Latin Anthesis ] takes place, when the burnt Anthers scatter their bags of Dust upon the Stigma.

Bractea of the female flowers very much enlarged after anthesis , when the spike presents the appearance of a pine-apple; bright yellow, with red apices.

The term anthesis is sometimes used to indicate the period at which the flower-bud opens.

There were both delayed and extended antheses and most of the time the flowers were semi-open.

Histologically the ovary and style are relatively simple at anthesis .

From the time of anthesis , when the floral parts open to receive pollen, the developing grain becomes the dominant sink.

A later planting date reduced pre-anthesis moisture stress by reducing the number of days..for the crop to reach anthesis .

• efflorescence 1626– The process of producing flowers, or bursting into flower; the period of flowering.
• blow 1748– Manner, style, or time of blossoming. Also figurative .
• anthesis 1783– The stage at which a flower is open, allowing fertilization to occur. Also: an instance of this.
• florescence 1793– The process of producing flowers or bursting into flower; the period or state of flowering. Also concrete . Flowers collectively.

Pronunciation

Plural: antheses.

• ð th ee
• ɬ rhingy ll

Some consonants can take the function of the vowel in unstressed syllables. Where necessary, a syllabic marker diacritic is used, hence <petal> /ˈpɛtl/ but <petally> /ˈpɛtl̩i/.

• a trap, bath
• ɑː start, palm, bath
• ɔː thought, force
• ᵻ (/ɪ/-/ə/)
• ᵿ (/ʊ/-/ə/)

Other symbols

• The symbol ˈ at the beginning of a syllable indicates that that syllable is pronounced with primary stress.
• The symbol ˌ at the beginning of a syllable indicates that that syllable is pronounced with secondary stress.
• Round brackets ( ) in a transcription indicate that the symbol within the brackets is optional.

View the pronunciation model here .

* /d/ also represents a 'tapped' /t/ as in <bitter>

Some consonants can take the function of the vowel in unstressed syllables. Where necessary, a syllabic marker diacritic is used, hence <petal> /ˈpɛd(ə)l/ but <petally> /ˈpɛdl̩i/.

• i fleece, happ y
• æ trap, bath
• ɑ lot, palm, cloth, thought
• ɔ cloth, thought
• ɔr north, force
• ə strut, comm a
• ər nurse, lett er
• ɛ(ə)r square
• æ̃ sal on

Simple Text Respell

Simple text respell breaks words into syllables, separated by a hyphen. The syllable which carries the primary stress is written in capital letters. This key covers both British and U.S. English Simple Text Respell.

b, d, f, h, k, l, m, n, p, r, s, t, v, w and z have their standard English values

• arr carry (British only)
• a(ng) gratin
• o lot (British only)
• orr sorry (British only)
• o(ng) salon

Inflections

anthesis typically occurs about 0.2 times per million words in modern written English.

anthesis is in frequency band 4, which contains words occurring between 0.1 and 1 times per million words in modern written English. More about OED's frequency bands

Frequency of anthesis, n. , 1810–2010

* Occurrences per million words in written English

Historical frequency series are derived from Google Books Ngrams (version 2), a data set based on the Google Books corpus of several million books printed in English between 1500 and 2010.

The overall frequency for a given word is calculated by summing frequencies for the main form of the word, any plural or inflected forms, and any major spelling variations.

For sets of homographs (distinct entries that share the same word-form, e.g. mole , n.¹, mole , n.², mole , n.³, etc.), we have estimated the frequency of each homograph entry as a fraction of the total Ngrams frequency for the word-form. This may result in inaccuracies.

Smoothing has been applied to series for lower-frequency words, using a moving-average algorithm. This reduces short-term fluctuations, which may be produced by variability in the content of the Google Books corpus.

Frequency of anthesis, n. , 2017–2023

Modern frequency series are derived from a corpus of 20 billion words, covering the period from 2017 to the present. The corpus is mainly compiled from online news sources, and covers all major varieties of World English.

Smoothing has been applied to series for lower-frequency words, using a moving-average algorithm. This reduces short-term fluctuations, which may be produced by variability in the content of the corpus.

Compounds & derived words

• synanthesis , n. 1880– Simultaneous ripening of the stamens and pistils in a flower.

Entry history for anthesis, n.

anthesis, n. was revised in March 2016.

anthesis, n. was last modified in July 2023.

oed.com is a living text, updated every three months. Modifications may include:

• further revisions to definitions, pronunciation, etymology, headwords, variant spellings, quotations, and dates;
• new senses, phrases, and quotations.

Revisions and additions of this kind were last incorporated into anthesis, n. in July 2023.

Earlier versions of this entry were published in:

OED First Edition (1885)

• Find out more

OED Second Edition (1989)

• View anthesis in OED Second Edition

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Citation details

Factsheet for anthesis, n., browse entry.

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Reduced nitrogen rate improves post-anthesis assimilates to grain and ameliorates grain-filling characteristics of winter wheat in dry land

• Research Article
• Published: 12 September 2023
• Volume 499 , pages 91–112, ( 2024 )

Cite this article

• Jinjin Wang   ORCID: orcid.org/0000-0002-3608-1856 1 , 2 , 3 ,
• Xu Sun 1 , 2 , 3   na1 ,
• Sadam Hussain 1 , 2 , 3 ,
• Lihua Yang 1 , 2 , 3 ,
• Sisi Gao 1 , 2 , 3 ,
• Peng Zhang 1 , 2 , 3 ,
• Xiaoli Chen 1 , 2 , 3 &
• Xiaolong Ren 1 , 2 , 3  

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Nitrogen (N) fertilizer application greatly enhances grain yield by improving dry matter accumulation and grain filling in winter wheat. However, the regulation mechanism of N rates on dry matter accumulation, transportation, and grain filling in winter wheat under dry farming could be more precise.

Five N treatments viz. 0, 75, 150, 225, and 300 kg·N·ha −1 , designated as N0, N75, N150, N225, and N300, were tested in two growing seasons of 2017–18 and 2018–19.

Fertilization significantly increased dry matter accumulation and intensity and assimilated accumulation after anthesis and its contribution to grain yield. N fertilizer significantly improved the superior and inferior grain weights, with the highest at N225, and increased by 22.29% and 24.41% in 2017–18 and 22.06% and 12.68% in 2018–19, respectively, compared to N0 ( P  < 0.05). Moreover, N fertilization increased the initial grain-filling potential and mean filling rate, shortened the days achieving the maximal grain-filling rate. N fertilizer increased the filling rate in the three periods, shortening the duration of the first-middle period, prolonging the late period, and increasing the contribution of the mid-late filling period to grain weight. However, excessive N application led to a reduction in the filling rate for each period. It prolonged the duration in the mid-late periods, which was not conducive to further increases in grain weight.

Conclusions

The N225 can be a more suitable N fertilizer for improving the accumulation and transfer of dry matter, promoting grain-filling, and increasing wheat yield in dry farming in the Loess Plateau.

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Changes in Nitrogen-Related Performance Attributes of Winter Wheat Varieties Released Between 1950 and 2020 in Dryland Region of China

Optimizing nitrogen fertilizer application under reduced irrigation strategies for winter wheat of the north China plain

Nitrogen supply modulates nitrogen remobilization and nitrogen use of wheat under supplemental irrigation in the North China Plain

Data availability.

The data are available from the corresponding author on reasonable request.

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This study was supported by the Program of Shaanxi Province Key R&D Program (No. 2021NY-073 and No.2022NY-196), National Natural Science Foundation of China (No. 31871580 and No. 31871562), Ningxia Hui Autonomous Region Key R&D Program (No. 2019BBF03011). We are grateful to Junfeng Nie, Baoping Yang, Ruixia Ding and Hui Li for help with the experiments.

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College of Agronomy, Northwest A&F University, Yangling, 712100, Shaanxi Province, China

Jinjin Wang, Xu Sun, Sadam Hussain, Lihua Yang, Sisi Gao, Peng Zhang, Xiaoli Chen & Xiaolong Ren

Key Laboratory of Crop Physiology, Ecology and Tillage in Northwest Loess Plateau, Minister of Agriculture, Yangling, 712100, Shaanxi Province, China

Institute of Water Saving Agriculture in Arid Areas of China, Northwest A&F University, Yangling, 712100, Shaanxi Province, China

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Jinjin Wang: Conceptualization, Methodology, Investigation, Formal analysis, Writing—original draft, Writing—review & editing. Xu Sun: Conceptualization, Methodology, Investigation, Formal analysis, Writing—original draft, Writing—review & editing. Sadam Hussain: Investigation, Formal analysis, Writing—original draft, Writing—review & editing. Lihua Yang: Investigation, Formal analysis, Writing—original draft, Writing—review & editing. Sisi Gao: Investigation, Formal analysis, Writing—original draft, Writing—review & editing. Peng Zhang: Writing—review & editing. Xiaoli Chen: Conceptualization, Methodology, Writing—review & editing. Xiaolong Ren: Conceptualization, Methodology, Writing—review & editing.

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Wang, J., Sun, X., Hussain, S. et al. Reduced nitrogen rate improves post-anthesis assimilates to grain and ameliorates grain-filling characteristics of winter wheat in dry land. Plant Soil 499 , 91–112 (2024). https://doi.org/10.1007/s11104-023-06276-0

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ORIGINAL RESEARCH article

Parental drought-priming enhances tolerance to post-anthesis drought in offspring of wheat.

• National Technique Innovation Center for Regional Wheat Production, Key Laboratory of Crop Physiology and Ecology in Southern China, Ministry of Agriculture, National Engineering and Technology Center for Information Agriculture, Nanjing Agricultural University, Nanjing, China

Drought is the major abiotic stress that decreases plant water status, inhibits photosynthesis, induces oxidative stress, restricts growth and finally lead to the reduction of wheat yield. It has been proven that drought priming during vegetative growth stage could enhance tolerance to drought stress at grain filling in wheat. However, whether drought priming imposed at grain filling in parental plants could induce drought tolerance in the offspring is not known. In this study, drought priming was successively applied in the first, the second and the third generation of wheat to obtain the plants of T1 (primed for one generation), T2 (primed for two generations), T3 (primed for three generations). The differently primed plants were then subjected to drought stress during grain filling in the fourth generation. Under drought stress, the parentally primed (T1D, T2D, T3D) plants, disregarding the number of generations, showed higher grain yield, leaf photosynthetic rate and antioxidant capacity as well as lower O 2 • − release rate and contents of H 2 O 2 and MDA than the non-primed (T0D) plants, suggesting that drought priming induced the transgenerational stress tolerance to drought stress. Moreover, the parentally primed plants showed higher leaf water status, which may result from the higher contents of proline and glycine betaine, and higher activities of Δ1-pyrroline-5-carboxylate synthetase (P5CS) and betaine aldehyde dehydrogenase (BADH), compared with the non-primed plants under drought stress. In addition, there was no significant difference among three generations under drought, and the drought priming in parental generations did not affect the grain yield of the offspring plants under control condition. Collectively, the enhanced accumulation of proline and glycine betaine in the parentally primed plants could have played critical roles in parental priming induced tolerance to drought stress. This research provided a potential approach to improve drought tolerance of offspring plants by priming parental plants.

Introduction

Drought is one of the critical environmental adversities affecting the growth, development and final yield of crop species ( Geng et al., 2016 ; Daryanto et al., 2017 ), and the frequency and severity of drought stress events are expecting to increase due to global climate change ( Cook et al., 2014 ; Zhao and Dai, 2015 ; Joshi et al., 2016 ). Drought stress perturbs a broad range of plant physiological and biochemical processes, including decreased plant water status, inhibited photosynthetic processes, induced oxidative stress damage and so on, which ultimately lead to growth retardation and the reduction of crop yield ( Perdomo et al., 2015 ; Saeidi and Abdoli, 2015 ; Daryanto et al., 2017 ). Wheat, one of the major food crop, is susceptible to drought stress especially during grain filling, which is the most critical stage determining the final grain weight ( Yang and Zhang, 2006 ). Therefore, improving plant tolerance to drought stress occurring during grain filling in wheat is meaningful for sustaining food security under the future climate.

Drought stress happened during grain filling could reduce wheat yield significantly, while drought tolerant varieties could maintain lower grain yield loss than the drought sensitive varieties ( Saeedipour and Moradi, 2011 ). Such yield loss was largely due to the inhibition of photosynthesis under drought ( Austin et al., 1977 ; Huseynova et al., 2016 ). The inhibition of plant photosynthesis may be resulted from the stomatal and/or non-stomatal limitation, which depended on the severity of drought stress ( Caemmerer and Farquhar, 1981 ; Lawlor and Tezara, 2009 ). In addition, the reactive oxygen species (ROS) burst under severe drought could cause oxidative stress damage, resulting in disturbance of a series of physiological and biochemical processes ( Cruz de Carvalho, 2008 ). Morphologically, plants under drought reduce the leaf area, decrease the stomatal conductance to reduce the water transpiration, and increase the root distribution in the deeper soil to maximum the water uptake ( Farooq et al., 2009 ; Fang et al., 2017 ). Besides, plant has developed a series of strategies at physiological, biochemical, and molecular levels to cope with drought stress, such as ABA content increase, ROS removement, osmotic adjustment and gene expression ( Chaves et al., 2003 ). The activation of antioxidant systems, which including enzymatic and non-enzymatic antioxidants, to remove excess ROS is an important strategy to cope with drought stress in plants ( Apel and Hirt, 2004 ). Antioxidant enzymes mainly include superoxide dismutase (SOD), catalase (CAT), ascorbate peroxidase (APX) and glutathione peroxidase (GPX), and non-enzymatic antioxidants mainly include reduced ascorbate (AsA) and reduced glutathione (GSH) ( Apel and Hirt, 2004 ; Miller et al., 2010 ). Another important biochemical response to drought is osmotic adjustment such as the accumulation of proline and glycine betaine (GB), which can help plants to retain or absorb more water by decreasing the osmotic potential of plant cells, buffer cellular redox potential and maintain the structure and physiological function of biological macromolecules ( Hare et al., 1998 ; Sakamoto and Murata, 2002 ; Ashraf and Foolad, 2007 ; Szabados and Savoure, 2010 ; Wang et al., 2014a ).

Several studies have shown that priming (pre-exposure to a moderate stress) could enhance tolerance to subsequent stresses, which is known as the term of “stress memory” ( Bruce et al., 2007 ; Chinnusamy and Zhu, 2009 ; Slaughter et al., 2012 ; Pastor et al., 2013 ). The stress-induced signaling chemicals, proteins, RNAs and metabolites were considered as short term memory factors ( Chinnusamy and Zhu, 2009 ), while epigenetic modifications such as DNA methylation and histone modifications are potential mechanisms for long-term, and even transgenerational memory ( Chinnusamy and Zhu, 2009 ; Hauser et al., 2011 ; Sani et al., 2013 ; Migicovsky et al., 2014 ). Our previous studies have found that drought priming during vegetative growth stage could enhance tolerance to freezing at jointing stage ( Li et al., 2015 ), and drought or heat during grain filling in wheat ( Wang et al., 2014b , c ). In addition, the offspring of the drought primed plants enhanced stress tolerance to post-anthesis high-temperature stress in wheat through improved photosynthesis and induced anti-oxidation capacity ( Zhang et al., 2016 ). Nosalewicz et al. (2016) reported that the intense drought stress has transgenerational effects on root morphology and topology of offspring in spring barley, where the offspring of the drought primed plants showed relatively decreased shoot-to-root ratio and reduced thick roots number, compared to the non-primed progeny under drought stress. It has been found that drought priming in the parental rice plants could induce proline accumulation through up-regulating expression of the proline synthesis genes paralleling with greater DNA demethylation in the offspring plants under drought ( Zhang et al., 2013 ). However, whether drought priming responses exist in the offspring of primed plants in wheat remains unclear.

In this study, we performed drought priming during grain filling in parental plants for three successive generations, and then subjected the respective offspring plants to drought stress during grain filling. Grain yield, photosynthesis, antioxidant system and osmotic adjustment were analyzed to investigate whether parental drought-priming could enhance tolerance to post-anthesis drought in offspring. The following hypotheses were tested: (i) drought priming in parental plants could enhance tolerance to drought stress in the offspring of wheat; and (ii) osmolytes accumulation may play critical roles in drought priming induced transgenerational drought tolerance.

Materials and Methods

Experimental setup.

Drought priming was performed in the cement pools (4 m in length, 2.5 m in width and 0.6 m in depth) with rain-proof shelter as reported in our previous study ( Zhang et al., 2016 ) at the Experimental Station of Nanjing Agricultural University, Nanjing, Jiangsu Province, China. The winter wheat ( Triticum aestivum L.) cultivar Ningmai 13 was used. As shown in Figure 1 , the 7-day drought priming event was successively applied in the first, the second and the third growth season (experimental year) to obtain the plants of T1 (primed for one generation), T2 (primed for two generations), T3 (primed for three generations). The plants without drought priming were annotated as T0. Leaf relative water content (LRWC) after drought treatment and the effect of drought on kernel weight of the parental plants were measured (Supplementary Table 1).

FIGURE 1. Setup of the experiment. The 7-day drought priming events (NP- no priming, DP- drought priming) were successively applied in the 1st, 2nd, and 3rd growth season (experimental year), the 5-day drought stress (NS, no drought stress; DS, drought stress) was conducted in the 4th year. Both the drought priming and the drought stress were initiated at 10 days after anthesis. In total, eight treatments were given: T0C, no former-generation priming + no offspring drought stress; T0D, no former-generation priming + offspring drought stress; T1C, one-generation priming + no offspring drought stress; T1D, one-generation priming + offspring drought stress; T2C, two-generation priming + no offspring drought stress; T2D, two-generation priming + offspring drought stress; T3C, three-generation priming + no offspring drought stress; T3D, three-generation priming + offspring drought stress.

The 5-day drought stress was conducted in the fourth year in a rainproof shelter. T0, T1, T2, T3 seeds were grown in plastic pots (22 cm in height and 25 cm in diameter) filled with 7.5 kg of soil (clay soil:sand = 2:1, w/w) with 0.9 g N, 0.36 g P 2 O 5 , and 0.9 g K 2 O mixed per pot. Another 1.6 g N was provided at the jointing stage. Prior to drought treatment, the seedlings were grown with normal water supply. At 10 days after anthesis, half of the T0, T1, T2, T3 seedlings were subjected to drought stress (D) by withholding water until soil relative water content (SRWC) decreased to ca. 40%, and then kept for 5 days before re-watering to the normal level. The SRWC of the rest seedlings maintained at ca. 75% were corresponding control (C). SRWC was measured according to the method of Wang et al. (2014b) . Taken together, eight treatments were established as T0D, T0C; T1D, T1C; T2D, T2C; T3D, T3C (Figure 1 ). The experiment was a randomized complete block design, with 30 pots as replicates for each treatment. The flag leaves of each treatment were collected at the third day for gene expression analysis and fifth day for determination of physiological and biochemical indexes during drought stress.

Plants Water Status

Leaf relative water content (LRWC) of flag leaves was measured as described by Jensen et al. (2000) . Predawn water potential of flag leaves (Ψ w ) was measured with an L-51 leaf hygrometer using HR-33T dew point microvolt-meter (Wescor Inc., Logan, UT, United States). Leaf osmotic potential (Ψ s ) was determined using a vapor pressure osmometer (Wescor 5600, Wescor Inc., Logan, UT, United States) at 25°C.

Photosynthesis and Chlorophyll Fluorescence Parameters

Gas exchange of flag leaves was measured on the last day of drought stress using a LI-6400 portable photosynthesis measurement system (LI-COR Biosciences, Lincoln, NE, United States) between 9:00 am and 11:00 am. The CO 2 concentration in the leaf chamber was set at 380 μmol mol -1 and the photosynthetically active radiation (PAR) was set at 1000 μmol m -2 s -1 .

The maximum quantum efficiency of photosystem II (Fv/Fm) after full dark adaptation (30 min) and the actual photochemical efficiency (ΦPSII) were measured with a portable chlorophyll fluorometer PAM-2500 (Heinz Walz GmbH, Eichenring, Effeltrich, Germany). Photochemical quenching (qP) and non-photochemical quenching (NPQ) of chlorophyll fluorescence were calculated according to the description of Maxwell and Johnson (2000) .

Antioxidant System

Content of malondialdehyde (MDA), the final product of lipid peroxidation, was measured as described by Du and Bramlage (1992) . O 2 • − release rate was assayed following hydroxylamine method ( Elstner and Heupel, 1976 ), and H 2 O 2 content was determined according to the method of Sui et al. (2007) . Activities of superoxide dismutase (SOD; EC 1.15.1.1) and catalase (CAT; EC1.11.1.6) were measured according to our previous methods ( Tan et al., 2008 ). Activities of ascorbate peroxidase (APX; EC 1.11.1.11), dehydroascorbate reductase (DHAR; EC 1.8.5.1) and monodehydroascorbate reductase (MDHAR; EC 1.6.5.4) were assayed by the method of Fryer et al. (1998) . Activities of glutathione peroxidase (GPX; EC 1.11.1.7) and glutathione reductase (GR; EC 1.6.4.2) were measured using Total Glutathione Peroxidase Assay Kit and Glutathione Reductase Assay Kit from Beyotime institute of Biotechnology (Shanghai, China), respectively. The contents of reduced ascorbate (AsA) and reduced glutathione (GSH) were measured as described by Gossett et al. (1994) . The content of soluble protein was measured by the method of Bradford (1976) .

Contents of Proline and GB

Proline content was determined according to ninhydrin coloring method ( Bates et al., 1973 ). 0.1 g of finely ground dried flag leaves was homogenized in 5 ml of 3% aqueous sulfosalicylic acid, heated for 10 min in boiling water bath followed by centrifugation. Two ml of supernatant was reacted with 2 ml of glacial acetic acid and 2 ml of acid-ninhydrin (2.5 g ninhydrin dissolved in 60 ml glacial acetic acid and 40 ml 6 M phosphoric acid) in a test tube for 1 h in boiling water bath. The reaction was terminated in an ice bath. The reaction mixture was mixed with 4 ml toluene completely and then standing for 2 h. The upper layer was used for determining the proline content at 520 nm using a spectrophotometer (UV-1780, Shimadzu (Suzhou) Instruments Manufacturing, Co., Ltd., Suzhou, China).

Glycine betaine (GB) content was measured following the description of Khoshro et al. (2013) . Five ml of toluene-water mixture (0.05% toluene) mixed with 0.2 g of finely ground dried flag leaves was mechanically shaken for 24 h at 25°C and then was filtered. Half ml of the filtrate was taken, and 1 ml of 2 N HCl solution, 0.1 ml of potassium tri-iodide solution (100 ml of 1 N HCl containing 7.5 g of iodine and 10 g of potassium iodide) was added. The mixture was incubated in an ice water bath for 90 min. After shaking gently, 10 ml of 1, 2-dichloroethane (chilled at -10°C) was poured into it. Then by passing a continuous stream of air for 2 min, two layers were separated. The absorbance of the organic layer was measured at 365 nm to determine the GB content.

Activities of Key Enzymes Involving in Proline and GB Metabolism

Extraction of Δ1-pyrroline-5-carboxylate synthetase (P5CS) and proline dehydrogenase (PDH) from wheat flag leaves was conducted according to the description of Tripathi et al. (2013) . Half gram of flag leaves were homogenized in 5 ml of 50 mM Tris-HCl buffer (pH 7.4) containing 3 mM EDTANa 2 , 7 mM MgCl 2 , 0.6 mM KCl, 1 mM DTT and 5% (w/v) PVP on ice. Then the homogenate was centrifuged at 4°C for 30 min at 14,000 g . The supernatant was used for enzymes activities determination.

P5CS activity was measured according to the method of Tripathi et al. (2013) . The reaction mixture volume was 3 ml containing 100 mM Tris-HCl (pH 7.2), 25 mM MgCl 2 , 75 mM sodium glutamate, 5 mM ATP, and 0.2 ml of extract. Then 0.2 ml of 0.4 mM NADPH was added in to initiate the reaction. The decrease in absorbance at 340 nm due to the consumption of NADPH was monitored to calculate P5CS activity.

PDH activity was assayed following the protocol of Tripathi et al. (2013) . The total reaction mixture volume was 3.5 ml containing 2.8 ml of 100 mM Na 2 CO 3 -NaHCO 3 buffer (pH 10.3, contained 20 mM L -proline) and 0.5 ml of enzyme extract, 0.2 ml of 10 mM NAD was finally added to initiate the reaction. The change in absorbance at 340 nm was monitored for calculation of PDH activity.

Betaine aldehyde dehydrogenase (BADH) activity was assayed following the method of Weretilnyk and Hanson (1989) . One fifth gram of flag leaves was homogenized in 10 ml of 0.1 M Tricine-KOH buffer (pH 8.5) containing 1 mM EDTANa 2 , 2 mM DTT, and 0.6 M sucrose. The homogenate was centrifuged at 10,000 g for 10 min at 4°C and the supernatant was collected. Protein in the supernatant was isolated by solid ammonium sulfate and desalted by centrifugal filtration on Sephadex G-25 columns. Enzyme extract (0.05 ml) was added in 0.95 ml of 0.1 M Tris-HCl buffer (pH 8.0) containing 0.5 mM NAD + and 5 mM DTT, reacted at 30°C for 30 min. BADH activity was measured at 340 nm and represented by NADH production amount per minute.

RNA Extraction, cDNA Synthesis and Quantitative Real-Time PCR

Total RNA was extracted from 50 to 100 mg of flag leaves using RNAiso Plus reagent (Takara Bio, Japan). The concentration and purity of RNA extract solution were measured with the NanoDrop 2000 (Thermo Scientific, United States) and the integrity was confirmed by agarose gel electrophoresis (1.2%). The removal of residual genomic DNA and first-strand cDNA synthesis was performed using HiScript II Q RT SuperMix for qPCR (+gDNA wiper) (Vazyme Bio, China) according to the instruction.

Sequences and its source of all primers were listed in Supplementary Table 2. Quantitative Real-time PCR was performed using ChamQ SYBR qPCR Master Mix (Vazyme Bio, China) on CFX Connect Real-Time PCR Detection System (Bio-Rad, United States) with cycling parameter: 95°C for 30 s; 40 cycles of 95°C for 10 s, 60°C for 30 s. Melting curves were run after PCR cycles. The relative expression levels of genes were calculated according to the 2 -ΔΔ Ct method, using ADP-RF gene as the reference gene. Three biological repeats and three technical repeats were performed.

ΔΔCt = (Ct target gene – Ct reference gene) treatment – (Ct target gene – Ct reference gene)T0C

Statistical Analysis

All data presented is the mean ± SE of three independent measurements. Data collected were analyzed using One-way ANOVA by SPSS package Ver. 22.0 (SPSS Inc., Chicago, IL, United States). Duncan’s multiple range test was used to determine significance differences among treatments ( P < 0.05).

Grain Yield and Yield Components

Grain yield was significantly reduced by drought, which was ascribed to the decrease in 1000-kernal weigh rather than number of ears and grain number per ear (Table 1 ). However, 1000-kernal weight yield of T1D (10.26%), T2D (12.02%), T3D (11.14%) was significantly higher than T0D, while there was no significant difference among T1D, T2D, T3D. In addition, T0C, T1C, T2C, and T3C showed similar 1000-kernal weight and grain yield.

TABLE 1. Effects of parental drought-priming on grain yield and yield components in offspring under drought stress during grain filling in wheat.

Plants Water Status, Photosynthesis and Chlorophyll Fluorescence

Drought significantly decreased leaf relative water content (LRWC), predawn water potential (Ψ w ) and osmotic potential (Ψ s ) of flag leaves (Figure 2 ). LRWC of T1D, T2D, T3D plants was 7.31, 8.47, and 6.37% higher respectively than the T0D. Ψ w of T1D, T2D, T3D plants were significantly higher while Ψ s was lower than T0D. However, there was no significant difference in these traits among T1D, T2D, and T3D. In addition, there was no difference in water status of flag leaves among T0C, T1C, T2C, and T3C.

FIGURE 2. Effects of parental drought-priming on water potential (Ψ w ), osmotic potential (Ψ s ) and relative water content (LRWC) of flag leaves of offspring plants under drought stress during grain filling in wheat. T0C, no former-generation priming + no offspring drought stress; T0D, no former-generation priming + offspring drought stress; T1C, one-generation priming + no offspring drought stress; T1D, one-generation priming + offspring drought stress; T2C, two-generation priming + no offspring drought stress; T2D, two-generation priming + offspring drought stress; T3C, three-generation priming + no offspring drought stress; T3D, three-generation priming + offspring drought stress. Data are means ± SE ( n = 3). Different lowercase letters indicate the significant difference at p < 0.05 level.

There was no significant difference in photosynthesis and chlorophyll fluorescence parameters of flag leaves among T0C, T1C, T2C, and T3C (Figure 3 ). Drought stress significantly decreased net photosynthetic rate (Pn) of flag leaves and T1D (42.32%), T2D (44.82%), T3D (45.10%) plants showed significantly higher Pn as compared with T0D. Stomatal conductance (gs) and transpiration rate (Tr) were significantly higher in T1D, T2D, T3D than in T0D, while intercellular CO 2 concentration (Ci) was lower. In addition, the maximum quantum efficiency of photosystem II (Fv/Fm) and the actual photochemical efficiency (ΦPSII) were also decreased by drought significantly. T1D, T2D, T3D showed higher Fv/Fm (3.26, 2.98, and 3.02%, respectively) and ΦPSII (11.97, 11.90, and 10.38%, respectively) than T0D. Photochemical quenching of chlorophyll fluorescence (qP) was significantly higher in T1D, T2D, T3D than in T0D, while non-photochemical quenching of chlorophyll fluorescence (NPQ) was lower. Again, there were no significant differences in these parameters among T1D, T2D, and T3D.

FIGURE 3. Effects of parental drought-priming on photosynthesis and chlorophyll fluorescence parameters of flag leaves of offspring plants under drought stress during grain filling in wheat. T0C, no former-generation priming + no offspring drought stress; T0D, no former-generation priming + offspring drought stress; T1C, one-generation priming + no offspring drought stress; T1D, one-generation priming + offspring drought stress; T2C, two-generation priming + no offspring drought stress; T2D, two-generation priming + offspring drought stress; T3C, three-generation priming + no offspring drought stress; T3D, three-generation priming + offspring drought stress. Data are means ± SE ( n = 3). Different lowercase letters indicate the significant difference at p < 0.05 level.

Antioxidant System in Flag Leaves

Under drought stress, MDA content was significantly increased (Figure 4 ). However, MDA content was much lower in T1D (21.33%), T2D (23.61%), T3D (19.25%) than in T0D. The O 2 • − release rate and H 2 O 2 content of flag leaves were significantly increased under drought stress, while they were less affected by drought in the primed (T1D, T2D, T3D) plants than in the non-primed (T0D) plants (Figure 4 ).

FIGURE 4. Effects of parental drought-priming on MDA content, O 2 • − release rate, and H 2 O 2 content in flag leaves of offspring plants under drought stress during grain filling in wheat. T0C, no former-generation priming + no offspring drought stress; T0D, no former-generation priming + offspring drought stress; T1C, one-generation priming + no offspring drought stress; T1D, one-generation priming + offspring drought stress; T2C, two-generation priming + no offspring drought stress; T2D, two-generation priming + offspring drought stress; T3C, three-generation priming + no offspring drought stress; T3D, three-generation priming + offspring drought stress. Data are means ± SE ( n = 3). Different lowercase letters indicate the significant difference at p < 0.05 level.

Activities of antioxidant enzymes such as SOD, CAT and APX were increased significantly by drought stress and were much higher in the primed (T1D, T2D, T3D) plants than in the non-primed (T0D) plants (Table 2 ). In addition, there was no significant difference in GPX activity among the non-drought treatments and T0D, while it was much higher in the primed (T1D, T2D, T3D) plants under drought.

TABLE 2. Effects of parental drought-priming on function of antioxidant system in flag leaves of offspring plants under drought stress during grain filling in wheat.

As for non-enzymatic ROS scavengers, the contents of AsA and GSH increased under drought stress (Table 2 ), moreover, the primed (T1D, T2D, T3D) plants had relatively higher GSH content than the non-primed (T0D) plants. However, there was no significant difference in AsA content among the drought stressed plants and among the control plants (Table 2 ). Activities of GR and DHAR were increased while of MDHAR was decreased by drought, however, they were all higher in the primed (T1D, T2D, T3D) plants than in T0D (Table 2 ). In addition, there was no significant difference in the above-mentioned traits in antioxidant system among T0C, T1C, T2C, and T3C. Gene expression of antioxidant enzymes was also assayed in this study (Supplementary Figure 1), only APX gene expression was consistent with its activity.

Proline and GB Accumulation

Contents of proline and GB significantly increased under drought stress, and the primed (T1D, T2D, and T3D) plants showed higher contents of proline (61.95, 64.07, and 60.11%, respectively) and GB (25.45, 20.80, and 33.89%, respectively) than the non-primed (T0D) plants (Figure 5 ). There was no significant difference among T0C, T1C, T2C, and T3C.

FIGURE 5. Effects of parental drought-priming on contents of proline and glycine betaine in flag leaves of offspring plants under drought stress during grain filling in wheat. T0C, no former-generation priming + no offspring drought stress; T0D, no former-generation priming + offspring drought stress; T1C, one-generation priming + no offspring drought stress; T1D, one-generation priming + offspring drought stress; T2C, two-generation priming + no offspring drought stress; T2D, two-generation priming + offspring drought stress; T3C, three-generation priming + no offspring drought stress; T3D, three-generation priming + offspring drought stress. Data are means ± SE ( n = 3). Different lowercase letters indicate the significant difference at p < 0.05 level.

Activities of P5CS, PDH, and BADH were measured to reveal the mechanism of accumulations of proline and GB induced by priming (Figure 6 ). Activities of P5CS and BADH were significantly enhanced while PDH activity was decreased due to the drought stress. The primed (T1D, T2D, T3D) plants had relatively higher P5CS (26.54, 25.43, and 23.52%, respectively) and BADH (23.68, 18.21, and 19.73%, respectively) activities than those of the non-primed (T0D) plants under drought. There was no significant difference in PDH activity between the primed (T1D, T2D, T3D) plants and the non-primed (T0D) plants, neither among T0C, T1C, T2C, and T3C plants.

FIGURE 6. Effects of parental drought-priming on activity of key enzymes involving in proline and glycine betaine metabolism in flag leaves of offspring plants under drought stress during grain filling in wheat. T0C, no former-generation priming + no offspring drought stress; T0D, no former-generation priming + offspring drought stress; T1C, one-generation priming + no offspring drought stress; T1D, one-generation priming + offspring drought stress; T2C, two-generation priming + no offspring drought stress; T2D, two-generation priming + offspring drought stress; T3C, three-generation priming + no offspring drought stress; T3D, three-generation priming + offspring drought stress. Data are means ± SE ( n = 3). Different lowercase letters indicate the significant difference at p < 0.05 level.

Transgenerational memory, which means the stress events happened during parental generation can be remembered by the plants and pass on to the next generation, and which could help the progeny more effectively to cope with the subsequent stresses ( Nosalewicz et al., 2016 ; Wang et al., 2016b ). The mechanisms of transgenerational priming induced tolerance are quite complex, and were reported to be associated with multiple physiological and molecular mechanisms, including epigenetic modifications (chromatin modification, miRNA etc.) ( Chinnusamy and Zhu, 2009 ; Sani et al., 2013 ; Migicovsky et al., 2014 ), plant hormone regulation, signaling pathway activation, osmotic adjustment and so on ( Hauser et al., 2011 ; Luna et al., 2012 ; Zhang et al., 2013 ). It is not clear whether the drought priming during the former generations could increase drought tolerance in offspring of wheat, and whether the priming responses are different among several successive priming generations. In this study, we imposed wheat plants to three successive generation drought priming and then subjected the offspring plants to drought stress. The results suggested that drought priming during parental generation could induce drought tolerance of offspring in wheat, as exemplified by less reduction grain yield of the primed (T1D, T2D, T3D) plants compared with the non-primed (T0D) plants under drought, which may results from better maintenance of photosynthesis, greater antioxidant capacity and higher osmolytes accumulation in the primed plants.

Parental Generation Drought Priming Enhances the Osmolytes Accumulation to Maintain Plant Water Status in Offspring Under Drought

Leaf water potential (Ψ w ) and relative water content (LRWC) are important indicators of plant water status ( Flower and Ludlow, 1986 ). In this study, after 5 days drought stress, Ψ w and LRWC of flag leaves was much lower in relation to the non-stressed plants. However, the primed (T1D, T2D, T3D) plants showed relatively higher LRWC and Ψ w than the non-primed (T0D) plants. The results were in line with our previous studies that drought priming during vegetative growth stages facilitated wheat plants to maintain higher leaf water status than the non-primed plants under drought during grain filling ( Wang et al., 2014c ). This indicates that parental drought priming could also help to maintain plant water status in offspring exposure to drought stress. Therefore, it is suggested that drought priming in parental plants could enhance drought tolerance in progenies plants.

The higher leaf water status may be owing to the higher accumulation of osmolytes to lower the osmotic potential, and that maintained turgor pressure ( Chaves et al., 2003 ). Proline and GB are two major organic osmolytes and play important roles in plant abiotic stress resistance, such as facilitate plants to retain or absorb water by decreasing the osmotic potential in cells and protect the PSII complex from damage ( Ashraf and Foolad, 2007 ; Wang et al., 2014a ). Accumulations of proline and GB were stimulated under drought stress and were significantly higher in tolerant cultivars than sensitive cultivars in wheat ( Ashraf and Foolad, 2007 ; Khoshro et al., 2013 ). In this study, the primed (T1D, T2D, T3D) plants had higher proline and GB contents than the non-primed (T0D) plants, which was corresponded to significantly lower Ψ s in the primed (T1D, T2D, T3D) plants. This indicates that drought priming in parental plants could decrease Ψ s of flag leaves at least partially via the accumulation of proline and GB hereby maintain turgor pressure and higher LRWC in offspring suffering from drought stress.

Glutamate pathway is the predominant pathway of proline synthesis under osmotic stress in plants ( Delauney et al., 1993 ), in which Δ1-pyrroline-5-carboxylate synthetase (P5CS) reduces glutamate to glutamate-semialdehyde (GSA), GSA then spontaneously converted to pyrroline-5-carboxylate (P5C) which is further reduced to proline by P5C reductase (P5CR). P5CS is the rate limiting enzyme in the process ( Hu et al., 1992 ; Verbruggen et al., 1993 ). In proline catabolism, proline dehydrogenase (PDH) first convert proline to P5C and P5C then converted to glutamate by P5C dehydrogenase (P5CDH), and PDH could be down-regulated by dehydration stress ( Kiyosue et al., 1996 ; Szabados and Savoure, 2010 ). In drought stressed wheat plants, P5CS was significantly induced, in paralleling with increased activity of P5CS and decreased activity of PDH, which promoted the accumulation of proline ( Aprile et al., 2009 ; Khoshro et al., 2013 ; Jiang et al., 2014 ). Our results showed that, the activities of P5CS increased and PDH decreased, these could contribute to the accumulation of proline under drought stress. The primed (T1D, T2D, T3D) plants showed higher activity of P5CS, and non-significant difference of PDH activity compared with the non-primed (T0D) plants, indicating that higher proline content induced in the primed plants was mainly ascribed to the elevated P5CS activity rather than modification of PDH activity.

BADH (betaine aldehyde dehydrogenase) is one of the key enzymes for GB biosynthesis ( Sakamoto and Murata, 2002 ). BADH could be induced and showed positive correlation with the GB content under drought stress in wheat ( Khoshro et al., 2013 ). In this study, BADH activity was increased significantly under drought stress and it was higher in the primed (T1D, T2D, T3D) plants than in the non-primed (T0D) plants, which was consistent with the higher GB content in these plants.

Parental Generation Drought Priming Improves Photosynthesis Performance and Final Yield in Offspring Under Drought

It has been reported that photosynthesis during grain filling stage contributes approximately 70–90% of photo-assimilates to the final grain yield under the favorable conditions ( Austin et al., 1977 ; Bidinger et al., 1977 ), while post-anthesis photosynthates are highly susceptible to drought ( Huseynova et al., 2016 ). Drought stress significantly restricts photosynthesis through stomatal limitation or non-stomatal limitation ( Caemmerer and Farquhar, 1981 ; Lawlor and Tezara, 2009 ). In this study, gs significantly decreased and Ci increased by drought stress, suggested that decreased photosynthesis rate was caused by both stomatal and non-stomatal limitation ( Farquhar and Sharkey, 1982 ). However, the parentally primed (T1D, T2D, T3D) plants showed higher Pn and gs as well as lower Ci than T0D, suggesting that there was less inhibition of stomatal and non-stomatal factors in the primed plants. Fv/Fm, ΦPSII and qP, which can reflect the capacity of photosystem II, can be reduced by severe drought stress ( Maxwell and Johnson, 2000 ; Wang et al., 2016a ). Here, the higher Fv/Fm, ΦPSII and qP values as well as lower NPQ observed in the primed (T1D, T2D, T3D) plants indicates that parental drought priming contributed to maintain relatively higher potential quantum efficiency, electron transport rate and the actual photosynthetic ability of PSII in offspring of wheat under drought stress, which is in accordance with higher Pn in the primed plants.

Grain yield data in the present study was consistent with photosynthesis, and the impact of drought on grain yield was mainly ascribed to the decline of 1000-kernal weight since the drought stress was applied during the grain filling stage. 1000-kernal weight yield of the primed (T1D, T2D, T3D) plants was higher than the non-primed (T0D) plants. This is in line with priming effect during seedling and stem elongation in our previous study ( Wang et al., 2014b ). Since the post-anthesis photoassimilates contribute the most part of grain filling ( Austin et al., 1977 ; Bidinger et al., 1977 ), the higher grain weight and final yield of the parentally primed (T1D, T2D, T3D) plants under drought treatment at least could be partially explained by the higher maintenance of the higher photosynthesis capacity than the non-primed (T0D) plants.

Parental Generation Drought Priming Contributes to Alleviate Oxidative Stress Damage in Offspring Under Drought

The inhibition of photosynthesis could lead to a higher leakage of electrons to O 2 by the Mehler reaction facilitating the production of a large amount of ROS under drought in wheat ( Cruz de Carvalho, 2008 ). ROS would further cause lipid peroxidation, MDA production and ultimately result in cell damage and plant death ( Cruz de Carvalho, 2008 ). In this study, we observed that MDA content was increased significantly by drought stress. However, it was much lower in the primed (T1D, T2D, T3D) plants than in the non-primed (T0D) plants. In line with MDA content, O 2 • − release rate and H 2 O 2 content of flag leaves were significantly increased under drought, while they were also less produced in the primed (T1D, T2D, T3D) plants than in the non-primed (T0D) plants.

Superoxide dismutase can dismutate superoxide to H 2 O 2 , acting as the first line of defense against ROS. Subsequently, CAT, APX and GPX were activated to detoxify H 2 O 2 ( Apel and Hirt, 2004 ). Antioxidant enzymes increased significantly under drought stress in wheat and the better drought tolerance performance in the tolerant cultivar was related to its higher antioxidant enzymes activities than the sensitive cultivar ( Zhang and Kirkham, 1994 ; Sairam and Saxena, 2000 ). In this study, drought stress significantly increased activities of SOD, CAT and APX to cope with increased O 2 • − release rate and H 2 O 2 content. Moreover, they were all higher in the primed (T1D, T2D, T3D) plants than in the non-primed (T0D) plants. For GPX activity, there was no significant difference among the non-drought treatments and T0D, while it was much higher in the primed (T1D, T2D, T3D) plants under drought. For other enzymes involving in A-G cycle and GPX cycle, activities of MDHAR, DHAR, and GR were all higher in the primed (T1D, T2D, T3D) plants than in T0D. As for non-enzymatic ROS scavenging mechanisms, AsA and GSH could be oxidized by H 2 O 2 so as to degrade H 2 O 2 ( Apel and Hirt, 2004 ). In this study, the contents of AsA and GSH increased by drought stress, and the primed (T1D, T2D, T3D) plants showed higher GSH content than the non-primed (T0D) plants. There was no significant difference in AsA content under drought. The increased activities of enzymes and higher GSH content contributed to the lower O 2 • − release rate, H 2 O 2 content and MDA content of flag leaves in the primed (T1D, T2D, T3D) plants than in the non-primed (T0D) plants. The above results illustrate that drought priming in parental plants could induce up-regulation of the antioxidant defense system in both enzymatic and non-enzymatic approaches to alleviate oxidative damage in offspring plants.

Research has shown that the induction of transcripts that encode antioxidant enzymes plays critical roles in cellular redox homeostasis ( Dutilleul et al., 2003 ). Thus, gene expression of antioxidant enzymes was also measured in this study. Transcript levels of CAT, APX, GPX, GR, MDHAR and DHAR were all increased under drought stress, consistent with enzymes activities except MDHAR. However, only APX showed higher transcript levels in the primed (T1D, T2D, T3D) plants than in the non-primed (T0D) plants. Therefore, higher activities of antioxidant enzymes in the primed (T1D, T2D, T3D) plants may be results of other explanations rather than the up-regulation of their encoding genes.

It has been proved that proline and GB play multiple functions in plants under stress, such as stabilizes the redox balance in photosystem, scavenges ROS; protects protein stability and enhances the activities of antioxidant enzymes ( Chaves et al., 2003 ; Szabados and Savoure, 2010 ). In accordance with this, the increased accumulation of proline and GB in the primed (T1D, T2D, T3D) plants may contribute to protect PSII and alleviate damages from drought to PSII, resulting in higher photosynthetic capacity as well as less production of ROS. Moreover, enhanced proline production could scavenge more ROS and contribute to a higher maintenance of activities of antioxidant enzymes discussed above.

Ding et al. (2012) found that the transcription rate and transcript levels of a subset of dehydration-response genes were increased significantly during recurring dehydration stresses in the same generation in Arabidopsis thaliana , which was then defined as “transcriptional memory” and this kind of genes were named as “trainable genes.” Luna et al. (2012) reported that progeny from bacteria-inoculated Arabidopsis (P1) were primed to activate salicylic acid-inducible defense genes of which promoters had changed histone modifications and were more resistant to the bacteria infection reoccurring again. There are few researches to investigate the effects of drought stress priming other than dehydration stress on the stress tolerance of the next generation other than the same generation in wheat. In this study, the elevated activities of P5CS and BADH in the parentally primed (T1D, T2D, T3D) plants may be resulted from up-regulation of P5CS and BADH expression, P5CS and BADH may be “trainable genes” playing roles in the transgenerational drought stress memory, or they could be regulated by the upstream signals which related genes have been trained. The “memory” factors should be changes of epigenetic modifications such as DNA methylation and histone modifications in “trainable genes.” In addition, ABA plays important roles in priming induced drought stress tolerance ( Wang et al., 2014c ) and cold stress tolerance ( Li et al., 2015 ) in the same generation, and the roles and mechanisms of ABA in priming induced transgenerational stress tolerance are far from clear and need further study.

It is interesting that there was no difference in all of the traits measured in this study among the primed (T1C, T2C, T3C) plants and non-primed (T0C) plants under non-drought conditions. This suggested that the parentally drought primed plants did not affect offspring plants in the physiological level under non-drought conditions, which was consistent with our previous finding ( Wang et al., 2016b ; Zhang et al., 2016 ). Furthermore, there was no significant difference in most traits we measured among three generations of the parentally primed (T1D, T2D, T3D) plants under drought stress. Research in Arabidopsis thaliana has shown that transgenerational memory may be unstable and occur in a stochastic manner ( Pecinka et al., 2009 ; Lang-Mladek et al., 2010 ). Therefore the improvement in drought tolerance by priming among the three generations is not suggested as a simple additive effect.

In conclusion, drought priming in parental plants could induce transgenerational stress tolerance to drought in offspring. Under drought stress, the parentally primed plants elevated activities of P5CS and BADH, which contributed to the enhanced accumulation of proline and GB. The accumulation of proline and GB played critical roles in osmotic adjustment to maintain higher plant water status, and also may contribute to less inhibition of photosynthesis, and higher ROS scavenging capacity (Figure 7 ). Therefore, the enhanced accumulation of proline and GB could play critical roles in parental priming induced alleviation of the drought damages. There was no significant difference in the alleviation effects on drought stress induced by different generations of priming, implying that one generation’s priming is enough to improve the tolerance of offspring plants to drought stress. In addition, the parental drought priming had no significant effect on offspring in terms of physiological processes and grain yield under non-drought conditions. This infers a potential approach to cope with the unpredicted drought stress by parental abiotic stress priming without side effect if the drought stress does not occur.

FIGURE 7. Mechanisms of parental drought-priming enhances tolerance to post-anthesis drought in offspring of wheat. The abbreviations of the enzymes or processes in bold were those included in the study, those in red, green and black indicate being up-regulated, down-regulated and not affected by the priming and stress, respectively. The parental drought-priming induces up-regulation of P5CS and BADH activities, which promotes the accumulation of proline and GB in offspring plants under the post-anthesis drought stress. Proline contributes to maintain leaf water status, stabilize the redox balance in photosystem, and enhance the activities of antioxidant enzymes (SOD, CAT, and enzymes in A-G cycle and GPX cycle), so as to decrease the production of MDA and then protect protein stability. And enhanced accumulation of GB could alleviate damages to PSII through accelerating D1 protein turnover to alleviate the photo-damage under drought.

Author Contributions

DJ and XW designed the experiments. XlW, XZ, and JCh performed the experiments and data analysis. XlW, XW, QZ, JCa, TD, WC, and DJ involved in the results discussion, manuscript writing and revising. All authors have read and approved the final manuscript.

This study was supported by projects of the National Key Research and Development Program of China (2016YFD0300107), National Natural Science Foundation of China (31325020, 31401326, 31471445, and 31771693), the China Agriculture Research System (CARS-03), Jiangsu Collaborative Innovation Center for Modern Crop Production (JCIC-MCP), and the National Non-profit Program by the Ministry of Agriculture (201403039).

Conflict of Interest Statement

The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.

Supplementary Material

The Supplementary Material for this article can be found online at: https://www.frontiersin.org/articles/10.3389/fpls.2018.00261/full#supplementary-material

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Zhang, X., Wang, X., Zhong, J., Zhou, Q., Wang, X., Cai, J., et al. (2016). Drought priming induces thermo-tolerance to post-anthesis high-temperature in offspring of winter wheat. Environ. Exp. Bot. 127, 26–36. doi: 10.3389/fpls.2016.00501

Zhao, T. B., and Dai, A. G. (2015). The magnitude and causes of global drought changes in the twenty-first century under a low-moderate emissions scenario. J. Clim. 28, 4490–4512. doi: 10.1175/JCLI-D-14-00363.1

Keywords : drought priming, drought stress tolerance, photosynthesis, osmolytes, antioxidant capacity, winter wheat ( Triticum aestivum L.)

Citation: Wang X, Zhang X, Chen J, Wang X, Cai J, Zhou Q, Dai T, Cao W and Jiang D (2018) Parental Drought-Priming Enhances Tolerance to Post-anthesis Drought in Offspring of Wheat. Front. Plant Sci. 9:261. doi: 10.3389/fpls.2018.00261

Received: 23 November 2017; Accepted: 14 February 2018; Published: 01 March 2018.

Reviewed by:

Copyright © 2018 Wang, Zhang, Chen, Wang, Cai, Zhou, Dai, Cao and Jiang. This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY) . The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.

*Correspondence: Dong Jiang, [email protected] Xiao Wang, [email protected]

Disclaimer: All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article or claim that may be made by its manufacturer is not guaranteed or endorsed by the publisher.

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Definition of antithesis

Did you know.

Writers and speechmakers use the traditional pattern known as antithesis for its resounding effect; John Kennedy's famous "ask not what your country can do for you—ask what you can do for your country" is an example. But antithesis normally means simply "opposite". Thus, war is the antithesis of peace, wealth is the antithesis of poverty, and love is the antithesis of hate. Holding two antithetical ideas in one's head at the same time—for example, that you're the sole master of your fate but also the helpless victim of your terrible upbringing—is so common as to be almost normal.

Examples of antithesis in a Sentence

These examples are programmatically compiled from various online sources to illustrate current usage of the word 'antithesis.' Any opinions expressed in the examples do not represent those of Merriam-Webster or its editors. Send us feedback about these examples.

Word History

Late Latin, from Greek, literally, opposition, from antitithenai to oppose, from anti- + tithenai to set — more at do

1529, in the meaning defined at sense 1b(1)

Dictionary Entries Near antithesis

anti-theoretical

Cite this Entry

“Antithesis.” Merriam-Webster.com Dictionary , Merriam-Webster, https://www.merriam-webster.com/dictionary/antithesis. Accessed 31 May. 2024.

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Kids definition of antithesis, more from merriam-webster on antithesis.

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Antithesis Definition

What is antithesis? Here’s a quick and simple definition:

Antithesis is a figure of speech that juxtaposes two contrasting or opposing ideas, usually within parallel grammatical structures. For instance, Neil Armstrong used antithesis when he stepped onto the surface of the moon in 1969 and said, "That's one small step for a man, one giant leap for mankind." This is an example of antithesis because the two halves of the sentence mirror each other in grammatical structure, while together the two halves emphasize the incredible contrast between the individual experience of taking an ordinary step, and the extraordinary progress that Armstrong's step symbolized for the human race.

Some additional key details about antithesis:

• Antithesis works best when it is used in conjunction with parallelism (successive phrases that use the same grammatical structure), since the repetition of structure makes the contrast of the content of the phrases as clear as possible.
• The word "antithesis" has another meaning, which is to describe something as being the opposite of another thing. For example, "love is the antithesis of selfishness." This guide focuses only on antithesis as a literary device.
• The word antithesis has its origins in the Greek word antithenai , meaning "to oppose." The plural of antithesis is antitheses.

How to Pronounce Antithesis

Here's how to pronounce antithesis: an- tith -uh-sis

Antithesis and Parallelism

Often, but not always, antithesis works in tandem with parallelism . In parallelism, two components of a sentence (or pair of sentences) mirror one another by repeating grammatical elements. The following is a good example of both antithesis and parallelism:

To err is human , to forgive divine .

The two clauses of the sentence are parallel because each starts off with an infinitive verb and ends with an adjective ("human" and "divine"). The mirroring of these elements then works to emphasize the contrast in their content, particularly in the very strong opposite contrast between "human" and "divine."

Antithesis Without Parallelism

In most cases, antitheses involve parallel elements of the sentence—whether a pair of nouns, verbs, adjectives, or other grammar elements. However, it is also possible to have antithesis without such clear cut parallelism. In the Temptations Song "My Girl," the singer uses antithesis when he says:

"When it's cold outside , I've got the month of May ."

Here the sentence is clearly cut into two clauses on either side of the comma, and the contrasting elements are clear enough. However, strictly speaking there isn't true parallelism here because "cold outside" and "month of May" are different types of grammatical structures (an adjective phrase and a noun phrase, respectively).

Antithesis vs. Related Terms

Three literary terms that are often mistakenly used in the place of antithesis are juxtaposition , oxymoron , and foil . Each of these three terms does have to do with establishing a relationship of difference between two ideas or characters in a text, but beyond that there are significant differences between them.

Antithesis vs. Juxtaposition

In juxtaposition , two things or ideas are placed next to one another to draw attention to their differences or similarities. In juxtaposition, the pairing of two ideas is therefore not necessarily done to create a relationship of opposition or contradiction between them, as is the case with antithesis. So, while antithesis could be a type of juxtaposition, juxtaposition is not always antithesis.

Antithesis vs. Oxymoron

In an oxymoron , two seemingly contradictory words are placed together because their unlikely combination reveals a deeper truth. Some examples of oxymorons include:

• Sweet sorrow
• Cruel kindness
• Living dead

The focus of antithesis is opposites rather than contradictions . While the words involved in oxymorons seem like they don't belong together (until you give them deeper thought), the words or ideas of antithesis do feel like they belong together even as they contrast as opposites. Further, antitheses seldom function by placing the two words or ideas right next to one another, so antitheses are usually made up of more than two words (as in, "I'd rather be among the living than among the dead").

Antithesis vs. Foil

Some Internet sources use "antithesis" to describe an author's decision to create two characters in a story that are direct opposites of one another—for instance, the protagonist and antagonist . But the correct term for this kind of opposition is a foil : a person or thing in a work of literature that contrasts with another thing in order to call attention to its qualities. While the sentence "the hare was fast, and the tortoise was slow" is an example of antithesis, if we step back and look at the story as a whole, the better term to describe the relationship between the characters of the tortoise and the hare is "foil," as in, "The character of the hare is a foil of the tortoise."

Antithesis Examples

Antithesis in literature.

Below are examples of antithesis from some of English literature's most acclaimed writers — and a comic book!

Antithesis in Charles Dickens' A Tale of Two Cities

In the famous opening lines of A Tale of Two Cities , Dickens sets out a flowing list of antitheses punctuated by the repetition of the word "it was" at the beginning of each clause (which is itself an example of the figure of speech anaphora ). By building up this list of contrasts, Dickens sets the scene of the French Revolution that will serve as the setting of his tale by emphasizing the division and confusion of the era. The overwhelming accumulation of antitheses is also purposefully overdone; Dickens is using hyperbole to make fun of the "noisiest authorities" of the day and their exaggerated claims. The passage contains many examples of antithesis, each consisting of one pair of contrasting ideas that we've highlighted to make the structure clearer.

It was the best of times , it was the worst of times , it was the age of wisdom , it was the age of foolishness , it was the epoch of belief , it was the epoch of incredulity , it was the season of Light , it was the season of Darkness , it was the spring of hope , it was the winter of despair , we had everything before us, we had nothing before us, we were all going direct to Heaven , we were all going direct the other way —in short, the period was so far like the present period, that some of its noisiest authorities insisted on its being received, for good or for evil, in the superlative degree of comparison only.

Antithesis in John Milton's Paradise Lost

In this verse from Paradise Lost , Milton's anti-hero , Satan, claims he's happier as the king of Hell than he could ever have been as a servant in Heaven. He justifies his rebellion against God with this pithy phrase, and the antithesis drives home the double contrast between Hell and Heaven, and between ruling and serving.

Better to reign in Hell than serve in Heaven.

Antithesis in William Shakespeare's Othello

As the plot of Othello nears its climax , the antagonist of the play, Iago, pauses for a moment to acknowledge the significance of what is about to happen. Iago uses antithesis to contrast the two opposite potential outcomes of his villainous plot: either events will transpire in Iago's favor and he will come out on top, or his treachery will be discovered, ruining him.

This is the night That either makes me or fordoes me quite .

In this passage, the simple word "either" functions as a cue for the reader to expect some form of parallelism, because the "either" signals that a contrast between two things is coming.

Antithesis in William Shakespeare's Hamlet

Shakespeare's plays are full of antithesis, and so is Hamlet's most well-known "To be or not to be" soliloquy . This excerpt of the soliloquy is a good example of an antithesis that is not limited to a single word or short phrase. The first instance of antithesis here, where Hamlet announces the guiding question (" to be or not to be ") is followed by an elaboration of each idea ("to be" and "not to be") into metaphors that then form their own antithesis. Both instances of antithesis hinge on an " or " that divides the two contrasting options.

To be or not to be , that is the question: Whether 'tis nobler in the mind to suffer The slings and arrows of outrageous fortune Or to take arms against a sea of troubles, And by opposing end them ...

Antithesis in T.S. Eliot's "Four Quartets"

In this excerpt from his poem "Four Quartets," T.S. Eliot uses antithesis to describe the cycle of life, which is continuously passing from beginning to end, from rise to fall, and from old to new.

In my beginning is my end . In succession Houses rise and fall , crumble, are extended, Are removed, destroyed, restored, or in their place Is an open field, or a factory, or a by-pass. Old stone to new building , old timber to new fires ...

Antithesis in Green Lantern's Oath

Comic book writers know the power of antithesis too! In this catchy oath, Green Lantern uses antithesis to emphasize that his mission to defeat evil will endure no matter the conditions.

In brightest day , in blackest night , No evil shall escape my sight. Let those who worship evil's might Beware my power—Green lantern's light!

While most instances of antithesis are built around an "or" that signals the contrast between the two parts of the sentence, the Green Lantern oath works a bit differently. It's built around an implied "and" (to be technical, that first line of the oath is an asyndeton that replaces the "and" with a comma), because members of the Green Lantern corps are expressing their willingness to fight evil in all places, even very opposite environments.

Antithesis in Speeches

Many well-known speeches contain examples of antithesis. Speakers use antithesis to drive home the stakes of what they are saying, sometimes by contrasting two distinct visions of the future.

Antithesis in Patrick Henry's Speech to the Second Virginia Convention, 1775

This speech by famous American patriot Patrick Henry includes one of the most memorable and oft-quoted phrases from the era of the American Revolution. Here, Henry uses antithesis to emphasize just how highly he prizes liberty, and how deadly serious he is about his fight to achieve it.

Is life so dear, or peace so sweet, as to be purchased at the price of chains and slavery? Forbid it, Almighty God! I know not what course others may take: but as for me, give me liberty or give me death .

Antithesis in Martin Luther King Jr.'s Oberlin Commencement Address

In this speech by one of America's most well-known orators, antithesis allows Martin Luther King Jr. to highlight the contrast between two visions of the future; in the first vision, humans rise above their differences to cooperate with one another, while in the other humanity is doomed by infighting and division.

We must all learn to live together as brothers —or we will all perish together as fools .

Antithesis in Songs

In songs, contrasting two opposite ideas using antithesis can heighten the dramatic tension of a difficult decision, or express the singer's intense emotion—but whatever the context, antithesis is a useful tool for songwriters mainly because opposites are always easy to remember, so lyrics that use antithesis tend to stick in the head.

Antithesis in "Should I Stay or Should I Go" by The Clash (1981)

In this song by The Clash, the speaker is caught at a crossroads between two choices, and antithesis serves as the perfect tool to express just how confused and conflicted he is. The rhetorical question —whether to stay or to go—presents two opposing options, and the contrast between his lover's mood from one day (when everything is "fine") to the next (when it's all "black") explains the difficulty of his choice.

One day it's fine and next it's black So if you want me off your back Well, come on and let me know Should I stay or should I go ? Should I stay or should I go now? Should I stay or should I go now? If I go, there will be trouble If I stay it will be double ...

Antithesis in "My Girl" by the Temptations (1965)

In this song, the singer uses a pair of metaphors to describe the feeling of joy that his lover brings him. This joy is expressed through antithesis, since the singer uses the miserable weather of a cloudy, cold day as the setting for the sunshine-filled month of May that "his girl" makes him feel inside, emphasizing the power of his emotions by contrasting them with the bleak weather.

I've got sunshine on a cloudy day When it's cold outside I've got the month of May Well I guess you'd say, What can make me feel this way? My girl, my girl, my girl Talkin' bout my girl.

Why Do Writers Use Antithesis?

Fundamentally, writers of all types use antithesis for its ability to create a clear contrast. This contrast can serve a number of purposes, as shown in the examples above. It can:

• Present a stark choice between two alternatives.
• Convey magnitude or range (i.e. "in brightest day, in darkest night" or "from the highest mountain, to the deepest valley").
• Express strong emotions.
• Create a relationship of opposition between two separate ideas.
• Accentuate the qualities and characteristics of one thing by placing it in opposition to another.

Whatever the case, antithesis almost always has the added benefit of making language more memorable to listeners and readers. The use of parallelism and other simple grammatical constructions like "either/or" help to establish opposition between concepts—and opposites have a way of sticking in the memory.

Other Helpful Antithesis Resources

• The Wikipedia page on Antithesis : A useful summary with associated examples, along with an extensive account of antithesis in the Gospel of Matthew.
• Sound bites from history : A list of examples of antithesis in famous political speeches from United States history — with audio clips!
• A blog post on antithesis : This quick rundown of antithesis focuses on a quote you may know from Muhammad Ali's philosophy of boxing: "Float like a butterfly, sting like a bee."

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• Open access
• Published: 30 January 2023

Timely sown maize hybrids improve the post-anthesis dry matter accumulation, nutrient acquisition and crop productivity

• R. R. Zhiipao 1 ,
• Vijay Pooniya 1 ,
• Niraj Biswakarma 1 ,
• Dinesh Kumar 1 ,
• Y. S. Shivay 1 ,
• Anchal Dass 1 ,
• Ganapati Mukri 2 ,
• K. K. Lakhena 1 ,
• R. K. Pandey 3 ,
• Arti Bhatia 4 ,
• Prabhu Govindasamy 1 ,
• Anamika Burman 1 ,
• Subhash Babu 1 ,
• R. D. Jat 5 ,
• A. K. Dhaka 5 &
• Karivaradharajan Swarnalakshmi 6  

Scientific Reports volume  13 , Article number:  1688 ( 2023 ) Cite this article

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• Agroecology
• Climate-change ecology
• Climate sciences
• Ecosystem ecology

Delayed sowing of maize hybrids could exacerbate the capability of maximizing the yield potential through poor crop stand, root proliferation, nutrient uptake, and dry matter accumulation coupled with the inadequate partitioning of the assimilates. This study appraised the performance of five recent maize hybrids viz., PMH-1, PJHM-1, AH-4158, AH-4271, and AH-8181 under timely and late sown conditions of the irrigated semi-arid ecologies. Timely sowing had the grain and stover yields advantage of 16–19% and 12–25%, respectively over the late sown maize hybrids. The advanced hybrids AH-4271 and AH-4158 had higher grain yields than the others. During the post-anthesis period, a greater dry matter accumulation and contribution to the grain yield to the tune of 16% and 10.2%, respectively, was observed under timely sown conditions. Furthermore, the nutrient acquisition and use efficiencies also improved under the timely sown. The nutrient and dry matter remobilization varied among the hybrids with AH-4271 and PMH-1 registering greater values. The grain yield stability index (0.85) was highest with AH-4158 apart from the least yield reduction (15.2%) and stress susceptibility index (0.81), while the maximum geometric mean productivity was recorded with the AH-4271 (5.46 Mg ha −1 ). The hybrids AH-4271 and PJHM-1 exhibited improved root morphological traits, such as root length, biomass, root length density, root volume at the V5 stage (20 days after sowing, DAS) and 50% flowering (53 DAS). It is thus evident that the timely sowing and appropriate hybrids based on stress tolerance indices resulted in greater yields and better utilization of resources.

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Inbred varieties outperformed hybrid rice varieties under dense planting with reducing nitrogen

Introduction.

Maize ( Zea mays L.), the third most cultivated crop after rice and wheat, can be grown in various soils and climates due to its versatility 1 , 2 . However, delayed sowing after an optimum time can result in reduced yields anomalies, due to aberrant weather events and irregular rainfall. Timely sowing is crucial for maximizing the yield of maize, and growers are concerned about the yield response of maize to sowing dates 3 . In addition, sowing at an optimum time could enhance the profitability of maize by improving the yields, as the crop has extended period to photosynthesize as well as avoid the artificial grain drying at the end of the crop cycle due to various environmental stresses 4 . The increased maize production is the result of improved agronomic management, varietal development 5 , and advances in plant protection measures. Furthermore, yield increment in a particular area is governed by the timely sowing, due to differences in climate and length of the growing season 6 . The supply of assimilate to grain in cereals derived from current assimilation, which supplies directly to kernels, and then remobilises the temporarily stored assimilates in the vegetative plant parts 7 , 8 . Reserve assimilates storage by a stem is strongly influenced by the growing conditions from emergence to the anthesis. Therefore, the grain yield could be buffered by the reserves accumulated in the vegetative portions of the plant pre-anthesis against the unfavourable current assimilation, particularly during the grain filling 8 .

In rice, an extended period of grain filling is the main determinant of grain yield, as it leads to a higher accumulation of dry matter because of greater cumulative mean temperatures and greater solar radiation interception 9 , 10 . The translocation of dry matter stored in the vegetative plant parts pre-anthesis and accumulation of photosynthates post-anthesis determined the grain yield of rice 11 . The dry matter translocated to grains during grain filling from the accumulated dry matter pre-anthesis and accumulation between flowering to physiological maturity acts as a function of grain yield in maize 12 , 13 , and differs among crop species and nutrient inputs 14 . In addition, it has been reported that ~ 85% of total grain dry weight was derived from the photo-assimilation during the grain filling period in maize 11 . Thus, the timely sowing of the genotypes improved the accumulation of photo-assimilates and their remobilization post-anthesis to grains. Further, timely sowing of maize hybrids could results in better interception of the photosynthetically active radiation leading to improved growth and development of the crop.

The identification of genotypes tolerant to stress and non-stress environment have been reported in crops using the indices, such as, stress susceptibility index (SSI), tolerance index (TOL), yield stability index (YSI), and geometric mean productivity (GMP) 15 , 16 . The SSI distinguishes the genotypes showing a minimum reduction in yield under the stress against the non-stress condition 17 , but it fails to identify the genotypes with high yield and stress-tolerant 16 . Further, the TOL indicates that the higher the TOL value more is tolerant to stress resulting in a higher yield potential of the genotypes 18 . Similarly, YSI is yet another index for identifying the stability of genotypes based on the yield under the stress and non-stress environment 19 . The GMP is similar to STI, wherein it indicates that the genotypes with higher GMP could be selected for both the stress and non-stress environment 15 . The above indices for the selection of genotypes have been reported mostly for wheat and other cereals by inducing an environment susceptible to the stress. Therefore, identifying the maize genotypes with higher yield potential under both stress and non-stress (timely and delayed sowing) environments would be a more robust and efficient approach.

In modern high-yielding maize hybrids, nitrogen use efficiency (NUE) has a negative correlation with the grain N concentration (GNC), defined as the grain yield per unit of nitrogen present in soil and applied fertilizer 16 , 20 , 21 . The decline in GNC could be attributed to the leaves staying green, which depends on the enhanced N absorption from the soil post-anthesis and less remobilization from the vegetative plant parts 22 , 23 . While genotypes with greater NUE under varied conditions can be beneficial in terms of protecting the environment, this should be considered when developing new varieties / hybrids and when recommending fertilizers such as N 24 . Moreover, greater N accumulation post-anthesis has a positive correlation with grain yield in rice, indicating that the post-anthesis N-accumulation plays an essential role in expanding the grain yield 25 . Subsequently, adequate concentration of phosphorus (P) is imperative for maintaining a high photosynthesis rate and enhancing the dry matter accumulation 26 . In addition, the extended period of leaf photosynthesis enhanced the grain yield but the trade-off with N and P remobilization in leaves during the grain filling as it shortens leaf photosynthesis 27 . In contrast, potassium (K) differs from P and N as it functions in various enzyme activation, synthesis of protein, maintaining osmotic balance and soluble metabolites translocation within plant tissue 28 . The hidden half of the plant (root systems) greatly influenced the performance of the above-ground portion particularly the formation of grain. Thus, the genotype with better rooting pattern could enhance the acquisition of limited resources (water, nutrients, etc.) and improved the yielding potential of the crop. Further, the root system architecture differs among genotypes, and so too does the nutrients and water uptake and ultimately the yield. Moreover, genotypes with improved root growth parameters and proliferation proportionately partitioned the captured resources and enhanced the yields 29 , 30 .

High-yielding hybrids are a boon for growers, but they require higher inputs as well as timely monitoring of all management practices. Hence, it is necessary to evaluate the performance of advanced/recent hybrids under the optimum nutrient management with varying sowing dates to know the productivity potential. This field study investigates the productivity, translocation and accumulation of dry matter besides the nutrient use-efficiencies and stress tolerance indices under the timely and late sowing of maize hybrids, in irrigated semi-arid ecologies.

Harvest index, grain and stover yields

On an average, the timely sown genotypes recorded 5–10% higher number of cobs per ha over the delayed sowing. While, among the hybrids, advanced hybrid AH-4271 recorded significantly a greater number of cobs ha −1 . Harvest index (HI) did not differ significantly under different sowing times, but advanced hybrids AH-4158 and AH-4271 showed significantly higher HI. Timely sowing gave a significantly stover yield advantage of 12–25.9% over the late sowing. PMH-1 and PJHM-1 had 5.2–16.9% and 5–11% greater stover yields than other hybrids in 2020 and 2021, respectively. With timely sowing of hybrid maize, grain yield increased by 16.7–19.2% compared to late sown hybrids. Hybrid AH-4271 had the highest grain yield, being similar to AH 4158, but significantly greater than the other hybrids (Table 1 ).

Yield attributes

The grains cob −1 under the timely sown was higher in 2020, but similar to the late sown in 2021. The highest grains cob −1 was recorded with AH-4271. Further, heavier cobs were harvested in timely sown than in late sown crop, though being similar in 2021. Among hybrids, the PMH-1 recorded the maximum cob weight, being similar to the AH-4158 and AH-4271. The timely sown crop produced 22.7% and 8.2% higher grain weight cob −1 over the late sown crop during 2020 and 2021, respectively. While, the grain weight among the hybrids were comparable, except AH-8181 in 2020, but in 2021 PMH-1 recorded the highest weight over other hybrids (Table 2 ).

Accumulated dry matter, its translocation and contribution to the grain yields

In 2020 and 2021, the late sowing led to a 6.3–14.6% increase in the dry matter translocation before anthesis (Fig.  1 a). Pre DMT translocation was the greatest with AH-8181 and PJHM-1. The effectiveness of dry matter translocation (Pre DMTe) was also higher under late sown than the timely sown crop (Fig.  1 b). While AH-8181 had the greatest Pre DMTe in 2020, PJHM-1 and AH-8181 in 2021. On the other hand, hybrids sown at the right time accumulated 26.5% (2020) and 5.7% (2021) greater dry matter after anthesis (Post DMA) (Fig.  1 c). PMH-1 and AH-4271 recorded the highest Post DMAs. Post anthesis dry matter accumulation efficiency (Post DMAe) under timely sowing was 2.4–19.9% greater than the late sowing (Fig.  1 d). PMH-1 and AH-4271 had a greater Post DMAe than the other hybrids. In 2020, the contribution of pre-anthesis dry matter translocation to grain yield (Pre DMTg) was higher for the late sowing (Fig.  1 e), while in 2021, it was the same for both the sowing dates (Fig.  1 f). In both the years, AH-8181 gave maximum Pre DMTg. Post-anthesis dry matter accumulation contributed 5.6–14.7% more to grain yield under the timely sowing compared to the late sowing. Hybrids AH-4271 and PMH-1 had similar Post DMAg in 2020, while PMH-1 had the maximum in 2021.

Pre DMT- pre-anthesis dry matter translocation ( a ), Pre DMTe- pre-anthesis dry matter translocation efficiency ( b ); Post DMA- post-anthesis dry matter accumulation ( c ), Post DMAe- post-anthesis dry matter accumulated efficiency ( d ); Pre DMTg- pre-anthesis dry matter translocation and Post DMAg- post- anthesis dry matter accumulation contribution to grain yield ( e , f ). Within years, sowing time, and genotypes, different letters on the individual bars of a figure indicate significant difference ( p  ≤ 0.05).

Nutrient concentration in different plant parts at flowering and maturity

Under the timely sown conditions, leaf and stem –N concentration at flowering was 2.9% and 8.8% greater than in the late sown, while hybrids didn’t differ significantly for leaf -N (Table 3 ). However, AH-4148 and AH-4271 had greater stem –N. Contrary to the N, the late sown crop had greater leaf and stem -P than timely sown, whereas hybrids had similar stem –P. For leaf –P concentration, PMH-1 being similar to AH-4158 and AH-8181, but greater than PJHM-1 and AH-4271. Leaf -K concentration in timely sown maize was 15.8% greater than that in the late sown. AH-4271 had greater leaf -K than that of PMH-1 and AH-8181, being similar to PJHM-1 and AH-4158. At maturity, the leaf and stem –N under timely sown was 10.7% and 3.8% greater, respectively over the late sown. The late seeded crop had higher leaf –P concentration than the timely sown, and the hybrid AH-8181 accumulated greater leaf –P than the other hybrids. Similarly, the leaf –P concentration was the highest in AH-8181, being comparable to PMH-1 and PJHM-1. While grain P in the hybrids was similar apart from the AH-4158. When sown early, the K concentration (leaf, stem, and grain) in maize hybrids was greater than when sown late. PJHM-1 had the highest leaf and grain –K concentration, while AH-8181 greater stem –K concentration.

Nitrogen translocation and uptake

Pre-anthesis N translocation in late-sown crop was 2.3–6.4% lower than in the timely-sown crop (Fig.  2 a). In both the years, hybrid AH-4158 exhibited significantly a greater translocation than other hybrids and was comparable to the hybrid AH-8181. The translocation efficiency (Pre NT eff.) was higher with the late-sown to the tune of 8.1–8.6% (Fig.  2 c). The hybrid AH-8181 achieved the best efficiency among the hybrids. After anthesis under the late sown condition, the N uptake (Post Nup) was 6.3% greater; however, timely sown had 12% greater uptake in 2021 (Fig.  2 b). As for N uptake, PJHM-1 was comparable to the AH-4271 and AH-8181, but significantly different from the PMH-1 and AH-4158. In 2021, the uptake of N by AH-4271 was 9.3 and 12.4% greater than that of PMH-1 and AH-8181. The proportion of N uptake after anthesis to total N accumulation (Post NR) varied with years and sowing times (Fig.  2 d). In 2020, PJHM-1 had significantly a greater Post NR than the other hybrids, but in 2021 the hybrids didn’t differ significantly.

Pre NT- pre-anthesis Nitrogen translocation ( a ), Post NuP- post-anthesis Nitrogen uptake ( b ), Pre NT-eff.- pre-anthesis Nitrogen translocation efficiency ( c ), Post NR- ratio of post-anthesis N uptake to total N accumulation ( d ). Within years, sowing time, and genotypes, different letters on the individual bars of a figure indicate significant difference ( p  ≤ 0.05).

Phosphorus (P) translocation and uptake

Pre-anthesis P translocation under the late sown crop was significantly greater than timely sown crop (Fig.  3 a). PMH-1 and AH-4271 had significantly greater pre-anthesis translocation rates than the other hybrids. Under late sowing, the translocation efficiency (Pre PT efficiency) was greater than the timely sowing (Fig.  3 b). In comparison with AH-4158, AH-4271 had significantly a higher Pre PT efficiency. Under timely sown conditions, the P uptake after anthesis (Post Pup) was 8.9–48.9% greater than the late sown crop (Fig.  3 c). In 2020, PJHM-1 and AH-4271 were similar, but significantly greater than other hybrids, whereas in 2021, it was PMH-1 and AH-8181. Under timely sowing, total P accumulation (Post PR) increased by 16.9–49.5% compared to the late sowing (Fig.  3 d). However, among the hybrids, in 2020 Post PR was highest with AH-4271, and in 2021, the PJHM-1 and AH-8181 were comparable, but significantly more than the other hybrids.

Pre PT- pre-anthesis phosphorus translocation ( a ), Post PuP- post-anthesis phosphorus uptake ( b ), Pre PT-eff.- pre-anthesis Phosphorus translocation efficiency ( c ), Post PR- ratio of post-anthesis P uptake to total P accumulation ( d ). Within years, sowing time, and genotypes, different letters on the individual bars of a figure indicate significant difference ( p  ≤ 0.05).

Potassium (K) translocation and uptake

The K translocation (Pre KT), its uptake after anthesis (Post Kup), and the ratio of K uptake to total K accumulation (Post KR) under timely sown were significantly greater than the late sown (Fig.  4 a–d). Compared to the late sowing, timely sown had an increments of 2.9–8.9% (Pre KT), 33.7–43.2% (Post Kup), and 24.5–36% (Post KR), respectively. PJHM-1 had the highest Pre KT and Pre KT efficiency than the other hybrids. Post Kup for AH-4158, AH-4271 and PMH-1 had recorded greater values than other hybrids. In 2020, AH-4158 and AH-4271 had comparable Post KRs, but AH-4271 recorded significantly more than other hybrids, however in 2021, PMH-1 recorded a statistically higher Post KR.

Pre KT- pre-anthesis potassium translocation ( a ), Post KuP- post-anthesis potassium uptake ( b ), Pre KT-eff.- pre-anthesis potassium translocation efficiency ( c ), Post KR- ratio of post-anthesis K uptake to total K accumulation ( d ). Within years, sowing time, and genotypes, different letters on the individual bars of a figure indicate significant difference ( p  ≤ 0.05).

Contribution of nutrient translocation and uptake to the grain yields

Despite of varying sowing dates and years, N translocation before anthesis did not affect the grain yields. Both PMH-1 and AH-4158 showed greater contributions to the pre-anthesis N translocation than did AH-4271 (Fig.  5 a,b). With late sown conditions, pre-anthesis P translocation was 14.8–53.3% greater than the timely sown crop (Fig.  5 c,d). Pre-anthesis translocation was largely accounted by AH-4158, followed by PMH-1. A greater K translocation was recorded under the late-sown conditions (Fig.  5 e,f). Meanwhile, PJHM-1 outperformed the other hybrids; timely sown hybrids contributed 10.8% more N to the grain yield from post-anthesis uptake (Fig.  5 a). In 2021, AH-4271 increased the post-anthesis uptake of N by 9.7–16.3%. Also, timely sowing increased the P uptake by 49.4% and 10.3%, respectively in 2020 and 2021 (Fig.  5 c,d). P uptake contributions were higher for AH-4271 and AH-8181 in 2020, whereas in 2021 they were PJHM-1 and AH-8181. Through post-anthesis K uptake, timely sowing contributed 21.1–29.7% more to grain yield over the late sowing (Fig.  5 e,f). AH-4158, AH-4271 and PMH-1 contributed most to the post-anthesis K translocation compared to other hybrids.

Pre NTg- Pre-anthesis nitrogen translocation and Post NupG- Post-anthesis nitrogen uptake contribution to grain yield ( a , b ); Pre PTg- Pre-anthesis phosphorus translocation and Post PupG- Post-anthesis phosphorus uptake contribution to grain yield ( c , d ); Pre KTg- Pre-anthesis potassium translocation and Post KupG- Post-anthesis potassium uptake contribution to grain yield ( e , f ).

Nutrient uptake in the shoot (above-ground) and efficiencies

Compared to late sown, the timely sown crop had a greater uptake of shoot N by 15.6% (2020) and 29.3% (2021) (Table 4 ). The hybrid AH-4271 was most effective when it came to absorbing N in the shoot. PMH-1 and AH-8181 recorded the highest values for shoot P uptake. Further, timely sown hybrids had 21.8% and 32.2% greater shoot K uptake than the late sown hybrids. In both the years, hybrids PMH-1, AH-4271 and PJHM-1 showed a greater shoot K uptake. Timely sown maize had 15.8–29.5% greater N uptake efficiency (NupE) than the late sown maize. In both the years, AH-4271 was the most efficient hybrid in absorbing N. On the other hand, the timely sown crop had a greater P uptake efficiency (PupE) than the late sown crops. The P uptake efficiency (PupE) in hybrids was greater for PJHM-1 in 2020, but for PMH-1 and AH-8181 in 2021. The K uptake efficiency (KupE) under timely sown was 21.7–32% greater than the late sown crop. PMH-1 and PJHM-1 gave the maximum K uptake efficiency in both years.

Nutrient (N, P and K) use efficiencies

The N use efficiency (NUE, kg kg −1 ) under timely sown conditions was 16.9–19.9% greater than the late sown conditions (Fig.  6 a). Also, AH-4158 and AH-4271 recorded the greater NUE. Timely sowing had 17.6–19.8% greater P use efficiency (PUE kg kg −1 ). Hybrid AH-4271 had the highest PUE (Fig.  6 b). Under timely sowing, KUE (kg kg −1 ) increased by 17.6–20.2% than the late sown conditions. Again, AH-4271 had the greater KUE and being similar to the AH-4158 (Fig.  6 c).

NUE- nitrogen use efficiency ( a ), PUE- phosphorus use efficiency ( b ), KUE- potassium use ( c ). Within years, sowing time, and genotypes, different letters indicate significant difference ( p  ≤ 0.05).

Stress tolerance indices

In terms of grain and stover yields, maize hybrids exhibited a variable response to the varying stress tolerance indices (Tables 5 , 6 ). The maximum grain yield reduction of 25.4% was recorded with the hybrid AH-4271. The most stable hybrids for grain yield, however, were AH-4158 and AH-8181. Furthermore, these hybrids also exhibited the least grain stress susceptibility index (SSI). The advanced hybrids AH-4271 (5.52 t ha −1 ) and AH-4158 (5.36 t ha −1 ) had the highest grain mean productivity (MP) and grain geometric mean productivity (GMP), respectively. In addition, the highest stover yield reduction was recorded with the AH-8181 (25.9%) due to the late planting. In terms of stover yield stability index (YSI), AH-4158 and AH-8181 were comparatively more stable with varying sowing times. In contrast, the hybrids with the highest stress susceptibility were PMH-1 and AH-8181. Nevertheless, the MP (7.45 t ha −1 ) and GMP (7.37 t ha −1 ) were highest with the hybrid PMH-1.

Root system traits

The maize hybrids had a significant variation in the root system traits, such as, root length (RL), root biomass (RB), root length density (RLD), and specific-root length (SRL). PJHM-1 had the highest root length, RLD, and SRL at the 20 DAS, while the maximum root biomass and volume was recorded with the advanced hybrid AH-4271 (Fig.  7 ). At 50% flowering (53 DAS), AH-4271 had the maximum RL, RLD, and SRL (Fig.  8 ), however, PJHM-1 had greater RB compared to other hybrids.

Root morphological traits of five maize hybrids grown in PVC tubes under field condition at V 5 stage (20 DAS). Means followed by different letters on the individual bars/lines of a figure indicate significant difference ( p  ≤ 0.05).

Root morphological traits of five maize hybrids grown in PVC tubes under field condition at 50% flowering (53 DAS). Means followed by different letters on the individual bars/lines of a figure indicate significant difference ( p  ≤ 0.05).

Timely sowing of maize hybrids enhanced the yields through better crop stand, improved yield attributes coupled with higher post-anthesis accumulation of dry matter and nutrient uptake. The contribution of post-anthesis dry matter accumulation to grain yield under both the sowing dates was much higher than the contribution from dry matter translocation. Indeed, the contribution of dry matter translocation to grain yield under the late sown was more than the timely sown, thereby indicates the importance of stored assimilates before anthesis under stress environment. While, for nutrients (N and K), the contribution to grain yield from translocation was more compared to the uptake, but the reverse hold true for P. Dry matter translocation and accumulation, nutrients translocation and uptake, and their contribution to grain yields varied significantly among the hybrids. Further, hybrids with higher geometric mean productivity and tolerance index are more productive under the varying sowing dates. On an average, timely sowing had a 17.9% and 18.9% yield advantage for grain and stover yields, respectively (Table 1 ), which could be attributed to the better photosynthates partitioning within the plant, as the crop has an extended period of photosynthesis 4 , with more favourable weather conditions during the growth and development for a particular region 6 . In addition, timely sowing also enhances the synchronization of maximum green leaf area index and the peak solar radiation 31 , thereby, improved the intercepted photosynthesis rate and hence the crop development 32 , resulting in greater yield. Additionally, the higher yield of hybrids in 2021 could be attributed to higher rainfall and its better distribution, particularly during the reproductive stage (Suppl. Figure  1 ).

The delayed sowing had the negative impacts on yields, by reducing the kernel number and their weight 33 . Comparable findings for the reduced kernel number and weight have also been reported by 5 , wherein late sowing would not be able to establish a proper root system under stress conditions. Hence, the uptake and partitioning of water and nutrients under the late sowing couldn’t meet the crop requirement for proper growth and development during the reproductive stage, which might have led to the under developed kernel. Subsequently, delayed sowing could also reduce the number, size and activity of growing grains coupled with the decreased supply of assimilate to grains during the period of grain filling, hence the grain yield 31 . There was a strong correlation between kernel weight with temperature and solar radiation 34 , and kernel weight with the grain yield 31 . Further, the main cause of the reduction in grain yield under late sowing was the reduction in grain number 35 .

Post-anthesis dry matter accumulation under timely sown accounted for 62.5% (av. of 2 yrs.) of the grain yield, wherein it was 55.8% greater than the late sown crop. While, it ranged from 55.6 to 67.7% in 2020 and 48.1 to − 64.1% in 2021 among the hybrids (Fig.  1 e,f), if respiratory losses for maintenance and remobilization of pre-anthesis accumulated assimilate are not taken into consideration 27 . The assimilates for grain formation don’t come entirely from the current assimilation which are directly transferred to the kernels, but also from the remobilization of temporarily stored assimilates in different vegetative plant parts 12 . Accumulated dry matter after anthesis is the major source for grain filling 13 , and in this study, the timely sown accumulated 17.2% higher dry matter and contributed 10.5% greater to the grain yield (Fig.  1 c,d) over the late sown crop. This could in fact be associated with the congenial environment for growth and development 36 , particularly during the grain filling period. In addition, hybrids with greater post-anthesis dry matter accumulation had the positive effects on grain yield, though PMH-1 had relatively lower grain yield which could be ascribed to the lower number of cobs per unit area (Table 2 ).

The N, P, and K accumulated in vegetative organs of the crop before anthesis, remobilized for grain filling. However, unlike the dry matter, the N, P, and K uptake post-anthesis could not meet the requirement for grain development 27 . Indeed, this study outlined that, the larger amount of N and K uptake occurred before anthesis, while the reverse is true for P (Figs.  2 a,b, 4 a,b) irrespective of the sowing dates and hybrids. The N and P are incorporated into the leaves and along with K it takes part in photosynthesis, hence recycling and remobilization of the stored N, P, and K in the vegetative tissues pre-anthesis would affect the photosynthesis processes 27 , 28 . It has been reported in maize and wheat that a greater amount of N and P accumulated during pre-anthesis is remobilized and recycled under the N and P deficiency 37 , 38 .

In the present study, timely sown had higher post-anthesis uptake of N, P, and K (Figs.  2 b, 3 b, 4 b) over the late sown crop, which could be the result of better growth and development of both above and the below-ground due to congenial environment, particularly during the grain filling period. Similar results of greater uptake of N, P, and K during the post-anthesis was reported by 39 . Further, the greater uptake of N, P, and K during the post-anthesis implies that it had the priority to be used in grain formation as can be visualized with the greater yields (Table 1 ). The greater post-anthesis uptake of N, P, and K under the timely sown crop is used to prolong the stay-green period of leaves, consequently, promotes more grain formation resulting in better yields 22 , 23 , but the grain % N was lower compared to the late sown crop (Table 3 ). Similar results of higher post-anthesis nutrient uptake with lower grain % N have also been reported by 20 , 21 , 27 .

The improvement in yields of maize hybrids under variable environmental conditions is desirable, hence in the present study various stress indices were employed to find out the best performing hybrid under the timely and late sown conditions. The higher rate of mean productivity (MP) and geometric mean productivity (GMP) coupled with a lesser stress susceptibility index (SSI) indicated that the genotypes had greater stress tolerance with the enhanced yield potential 15 . The hybrid AH-4158 had the least yield reduction and SSI with greater YSI (Table 5 ), thereby, it had greater tolerance with the time of sowing. Subsequently, AH-4271 outperformed for grain MP and GMP though it was more susceptible to late sowing. This hybrid AH-4271 would be well suited under the timely sown with higher productivity potential. In addition, this hybrid had greater stability for the stover yield with different sowing dates (Table 6 ).

A positive correlation was reported between SSI and grain yield of wheat genotypes to identify the best performing varieties under stress conditions 40 . Further, classifying the genotypes based on MP and GMP were similar, and had positive relations with the grain yield under normal and stress conditions 15 , 41 . The MP, which is the average productivity of yield under stress and normal conditions 42 , and its greater value denotes a better performance of hybrids under the stress conditions, hence a good criterion for selecting a hybrid tolerant to stress. The studies of drought stress on maize hybrids yield reported that under normal and mild stress conditions, the GMP, MP, and stress tolerance index (STI) were important indices for identifying the best performing hybrids under the variable environments 43 . Further, a positive correlation of grain yield with MP and GMP under severe stress and normal conditions were observed, thereby helps in determining the drought-tolerant hybrids 44 .

In our experiment, a hybrid with greater MP, GMP, and YSI could be used for identifying the hybrids adaptability to different sowing dates (timely and late). In terms of grain yield, the AH-4271 and AH-4158 had greater MP and GMP under the variable sowing dates, indicating their superiority. The adoption of N efficient hybrids is a vital management strategy for enhancing the N use efficiency (NUE) 45 . The NUE is grain yield per unit available-N both from soil and through applied fertilizer 16 . In cereals, the NUE has been reported to be about 40% of the applied fertilizers 46 . In our study, the NUE ranged between 10–15 kg kg −1 considering the contribution from 0.0–0.30 m soil profile in addition to the applied N fertilizer (Fig.  6 a).

The greater NUE of AH-4271 and AH-4158 under the timely sown conditions could be attributed to the enhanced uptakes by their root coupled with the better assimilation and remobilization in the shoot 16 , 47 . Studies on wheat showed that NUE could be improved through optimization of the root system 48 , 49 . Furthermore, significantly greater P and K use efficiencies under timely sown (Fig.  6 b,c) might be the result of better growth and development with higher grain yield due to the congenial crop environment. Subsequently, differential growth and development habits of the hybrids in changing environments lead to variation in P and K use efficiencies. The greater P and K efficiencies with the AH-4271 and AH-4158 (Fig.  6 b,c), showed their responsiveness and superiority through better adaptation under the varied ecologies. The importance of growing nutrients use efficient genotypes has been emphasized, as it would reduce the excessive fertilizers input without yield penalty 16 , 49 .

In addition, the rooting traits varied among the hybrids (Figs.  7 , 8 ), where in the hybrid with better root length and biomass coupled with greater root length density produced better yields (Table 1 ). The genotypes with better root proliferation at the early stage might have led to the better crop establishment and used the available resources more efficiently and partitioned proportionately to different plant parts, resulting in the greater output. Indeed, the greater number of cobs per unit area was recorded with those hybrids having better root morphological traits, thereby indirectly implies more crop stand on a unit area. Genotype with a greater root growth and proliferation proportionately partitioned the captured resources and thus enhanced the yields 29 , 30 . Similarly, studies on wheat showed that root biomass was positively correlated to the number of grains spike −1 and yields 50 , so was the case in the present study, i.e., a higher number of grains cob −1 and grain yields related.

Conclusions

Timely sowing, a resource-saving practice plays a vital role in enhancing the yield potential of maize genotypes. Compared with late sown, timely sown yielded 16–19% more grain and produced 5–10% more cobs per hectare. During the grain filling period, post-anthesis dry matter accumulation is crucial, and it was significantly higher with the timely sowing compared to the late sowing. We observed maximum nutrient use efficiency, nutrient uptake, and nutrient contribution to grain yield under the timely sowing conditions. Advanced hybrids, AH-4271 and AH-4158, performed better in various parameters, viz., grain yield, nutrient uptake, and cobs per ha. Hybrid AH-4158 showed higher yield stability, a lower stress susceptibility index, and a lower percentage of grain yield reduction, indicating the enhanced capacity for flexibility in a variety of crop-growing conditions, together with hybrid AH-4271. Indeed, better root morphology correlates with the greater nutrient use efficiency, dry matter accumulation, and nutrient remobilization to produce higher yields. Therefore, assessing hybrids based on stress indices, nutrient remobilization, and grain yield could lead to identifying the best hybrids under the variable crop conditions.

Materials and methods

Experimental site and weather conditions.

A Fixed-site field experiment was conducted for 2 years at the ICAR-Indian Agricultural Research Institute, New Delhi, India, planting maize genotypes under the timely and late sown conditions during the rainy seasons of 2020 and 2021. The region falls under the Trans Indo-Gangetic plains with 28°38′ N latitude, 77°10′ E longitude, and 229 m amsl. The climate is semi-arid with hot summers and rainy in monsoon (July–September), with scattered rains in winter. There is an annual mean rainfall of 650 mm, and the mean maximum and minimum temperatures range from 20–40 °C to 4–28 °C, respectively. The 2 years weather observations (2020–21) recorded by the automated observatory in the adjacent experimental site are summarized in suppl. Figure  9 . Before preparatory tillage, the soil samples were collected randomly from 0.0–0.15 m to 0.15–0.30 m undisturbed soil depth. the samples were air-dried, ground, sieved through a 0.2 mm sieve, and stored in air-tight polyethene bags for further analysis of soil chemical properties, viz. soil pH (1:2.5, soil: water 51 ), KMnO 4 -oxidizable N 52 , NaHCO 3 extractable P 53 , NH 4 OAc exchangeable K 54 , and soil organic carbon (S OC ) 55 (Table 7 ).

Cultural operations, experimental design, and crop management

Pre-sowing irrigation was applied before preparatory tillage operations. The field was deep ploughed twice using a disc harrow (0.00–0.20 m depth) followed by planking with a rotavator/ cultivator twice and finally levelled. The experiment was laid out in a split-plot design with three replicates. Two sowing times (timely and late sown) were allocated to the main plots and five recently released maize hybrids (PMH-1, PJHM-1, AH-4158, AH-4271, and AH-8181) to sub-plots, with a sub-plot size of 20 m 2 (4 m × 5 m). In 2020, the crop was sown on 6th July (timely) and 27th July (late), while in 2021 it was sown on 3rd July (timely) and 24th July (late), respectively. The seeds were dibbled manually at a spacing of 0.75 m (row–row) × 0.20 m (plant–plant) in both the seasons. Earthing-up was done at the knee-high stage for better crop growth, prevent lodging and uniform distribution of irrigation water. Based on the critical growth stages and the rainfall received during the crop seasons, irrigation water was applied to a depth of 0.05 m. The recommended fertilizer application rate for maize was 150:26.2:49.59 kg NPK ha −1 . Nitrogen (N) was applied as urea (46% N), phosphorus (P) and potassium (K) through di-ammonium phosphate (46% P 2 O 5 ), and muriate of potash (60% K 2 O), respectively. At sowing, full doses of P, K, and 50% N were applied uniformly in all the plots, while the remaining 50% N was top-dressed in two equal splits at knee-high and tasseling stages. Weeds were controlled through broad-spectrum pre-emergence herbicide atrazine (50% WP, at 750 g a.i. ha −1 ) applied a day after sowing, followed by one hand weeding at 35–40 days after sowing (DAS). For insect-pests management, particularly fall armyworm ( Spodoptera frugiperda ), a systemic insecticide emamectin benzoate (50% SG, at 200 g ha −1 ) was first sprayed at 20 DAS, followed by need-based at 15–20 d intervals on their appearance. The insect-pests and disease management were carried out uniformly in all the plots based on the recommended practices.

Plant sampling and their laboratory analysis

Plant samples were collected at two stages of the crop growth (tasseling and maturity) at two different dates in each season. In each plot, three plants were randomly cut at the base and separated into leaf, stalks (stem + leaf sheaths + tassel), and leaf, stalks (stem + leaf sheaths + tassel + husk), cob and grain at maturity. The samples were placed in a perforated brown paper bag, air-dried for 48 h, and then oven-dried at 65 ± 2 °C to a constant weight. The dry matter (DM) values were used to determine the translocation, accumulation, and efficiency as per the equations described by (1) 11 , 21 , 56 . A Macro Wiley-mill having a 40-mesh sieve was used for grinding the plant samples, and appropriate amounts (0.5 g) of the ground samples were used to determine the total N concentration employing the modified Kjeldahl digestion process, total P by colored Vanado-molybdo-phosphoric acid procedure, and total K by flame photometer method 57 . The nutrient translocation, uptake, and efficiency were computed in accordance with the Eq. (2) 21 , 58 . The soil available nutrients in this study were determined to the depths of 0.00–0.30 m. Also, the above-ground nutrient uptake at maturity, uptake efficiencies were computed by using Eq. (3) 59 , 60 . The above-ground nutrient uptake was worked out by multiplying the nutrient concentration in stalks and grains with the biomass yields. To estimate the uptake and use efficiencies, the soil nutrients available to a depth of 0.30 m (0.00–0.15 m, 0.15–0.30 m) were used.

• Pre NT, Pre NTe, Post Nup, Post NR, Pre NTg, Post NupG- Nitrogen; Pre PT, Pre PTe, Post Pup, Post PR Pre PTg, Post PupG- Phosphorus; Pre KT, Pre KTe, Post Kup, Post KR, Pre KTg, Post KupG- Potassium.
• AGN, NupE, NUE—Nitrogen; AGP, PupE, PupE—Phosphorus; AGK, KupE, KUE—Potassium.

The maize genotypes were subjected to mathematical relationships on the basis of stover and grain yields for identifying the best performing genotype under normal and the delayed sowing conditions. The following formulae were used to find out the efficient genotypes 15 , 16 .

Percentage reduction of yield (YR; %) = (Y T  − Y L )/(Y T  × 100)

Mean productivity (MP) = (Y T  + T L )/2

Stress susceptible index (SSI) = (1 − Y T /Y L )/SI. Where Stress intensity (SI) was calculated as, SI = 1 − (X T /X L )

Geometric mean productivity (GMP) = √(Y T  × Y L )

Yield Stability Index (YSI) = Y T /Y L

Where, Y T and Y L are the yields of genotypes under timely and late sown, respectively. The X T and X L denote mean yield of all genotypes under timely and late sown conditions, respectively.

Yield attributes and yields

The physiological growth stages were marked when 50% of the plants attained their particular stage, such as, tasseling, silking, and physiological maturity. In the first season, maize was harvested on 18th and 27th October 2020, while in the second season on 16th and 27th October 2021, respectively for the timely and late sown conditions. The crop was harvested from the middle three rows (4 m × 2 m, 8 m 2 ) leaving two border rows on each side. First, the cobs were hand-picked, then the stover was cut from the ground surface. The harvested produce was sun-dried for 25–30 d to bring down the grain moisture from 20–22% to 14–15% for threshing and the yield measurement for grain and stover were done separately. Further, the yield attributes, such as, number of rows cob −1 , number of grains cob −1 , cob weight (g), grains weight cob −1 (g), and 100-grains weight (TW, g) were determined from the five randomly picked cobs.

The plants were grown in 25 mm PVC tube, 0.195 m diameter, 0.5 m and 1.0 m deep, transparent cylindrical polyethylene sheet placed on the inner PVC tube, filled with soil to a depth of 1.0 m. The PVC tubes were placed vertically on the steel stands in an open field condition. The soil (0.0–0.15 m depth) for the above purpose was collected from the experimental field of the current study. The 5 mm sieve was used to sieve out the debris and other undesirable materials before being packed into the PVC tubes. The soil in the tubes was manually watered carefully to saturation level so as to avoid the excess drainage (Pooniya et al. 30 ). The soil in the tubes was fertilized at sowing to the depth of 0.01 m with an equivalent to the recommended fertilizer rate for maize hybrids (150:26.2:49.59 kg NPK ha −1 ), in the form of urea, diammonium phosphate, and muriate of potash. The plants were hand watered based on the critical growth stages as per the requirement. Three seeds each of the five recent/advanced hybrids were manually dribbled to a depth of 0.05 m in each tube, during the first week of July 2019. Five days after germination, a single plant was maintained in each tube after carefully thinning out the remaining seedlings. The five hybrids were grown in a completely randomized block design with three replicates and harvested twice {V5 stage-20 DAS (Fig.  9 a) and 50% flowering- 53 DAS (Fig.  9 b)}. At first sampling (0.5 m depth tube), the shoots were first cut from the roots at the crown, while in the second (1.0 m depth tube), the shoots and stilt roots above the soil were cut before washing. The polyethylene sheet was gently pulled from the tubes, cut open and dipped in the still water for an hour, repeatedly washed the soil on a 2 mm sieve to produce a clean root sample 29 . The recovered roots were placed in the plastic bags and stored at 5 °C until the scanning of its morphological traits. The scanned images of the roots were analyzed with WinRHIZO professional software (LA2400, Regent instrument, Quebec, Canada) for recording the root morphological traits (total root length and root volume, in the present study). The scanned roots were further dried in a hot air oven at 65 ± 5 °C until obtaining the constant weight for root dry biomass. The root length density was computed by dividing the total root length with the soil volume (0.0149 m 3 —1st sampling and 0.0298 m 3 —2nd sampling). While the specific root length was calculated as the total root length divided by the root biomass 30 .

Destructively measured root growth pattern of the maize genotypes (PMH-1, PJHM-1, AH-8181, AH-4271, AH-4158) grown in PVC tubes under the field conditions at V5 stage (20 days after sowing, DAS) ( a ) and at 53 DAS ( b ).

Statistical analysis

With SAS 9.4, the data were statistically analyzed for split-plot designs through analysis of variance (ANOVA) 61 . In addition, Tukey’s honestly significant difference at 0.05 probability ( p  ≤ 0.05) was used to compare the mean effects of the treatments i.e., sowing dates and genotypes.

Authors have confirmed that all the plant studies were carried out in accordance with relevant national, international or institutional guidelines.

Data availability

The data that support the findings of this study are available on request from the corresponding author. The data are not publicly available due to private and ethical restrictions.

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Acknowledgements

We acknowledge to the ICAR-Indian Agricultural Research Institute and the Indian Council of Agricultural Research, New Delhi for providing the necessary field and laboratory facilities during the experimentation.

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Agronomy, ICAR-Indian Agricultural Research Institute (IARI), New Delhi, 110 012, India

R. R. Zhiipao, Vijay Pooniya, Niraj Biswakarma, Dinesh Kumar, Y. S. Shivay, Anchal Dass, K. K. Lakhena, Prabhu Govindasamy, Anamika Burman & Subhash Babu

Genetics, ICAR-Indian Agricultural Research Institute (IARI), New Delhi, 110 012, India

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Plant Physiology, ICAR-Indian Agricultural Research Institute (IARI), New Delhi, 110 012, India

R. K. Pandey

Environment Science, ICAR-Indian Agricultural Research Institute (IARI), New Delhi, 110 012, India

Arti Bhatia

Chaudhary Charan Singh Haryana Agricultural University (CCSHAU), Hisar, Haryana, 125004, India

R. D. Jat & A. K. Dhaka

Microbiology, ICAR-Indian Agricultural Research Institute (IARI), New Delhi, 110 012, India

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R.R.Z., N.B., V.P., led the research work, planned, supervised, and conducted field experiments, and read and edited the manuscript. N.B., R.R.Z., K.K. L., A.B., collected soil/plant samples and performed chemical analysis, also wrote the initial draft of the manuscript, and prepared figures, and tables. D.K. Y.S., A.D., G.M., R.K.P., P.G., S.B., A.B., K.S., project supervision, reviewed, read and edited the manuscript with significant contributions. R.D.J. and A.K.D. performed the statistical analysis and prepared figures.

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Zhiipao, R.R., Pooniya, V., Biswakarma, N. et al. Timely sown maize hybrids improve the post-anthesis dry matter accumulation, nutrient acquisition and crop productivity. Sci Rep 13 , 1688 (2023). https://doi.org/10.1038/s41598-023-28224-9

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Timely sown maize hybrids improve the post-anthesis dry matter accumulation, nutrient acquisition and crop productivity

R. r. zhiipao.

1 Agronomy, ICAR-Indian Agricultural Research Institute (IARI), New Delhi, 110 012 India

Vijay Pooniya

Niraj biswakarma, dinesh kumar, y. s. shivay, anchal dass, ganapati mukri.

2 Genetics, ICAR-Indian Agricultural Research Institute (IARI), New Delhi, 110 012 India

K. K. Lakhena

R. k. pandey.

3 Plant Physiology, ICAR-Indian Agricultural Research Institute (IARI), New Delhi, 110 012 India

Arti Bhatia

4 Environment Science, ICAR-Indian Agricultural Research Institute (IARI), New Delhi, 110 012 India

Prabhu Govindasamy

Anamika burman, subhash babu.

5 Chaudhary Charan Singh Haryana Agricultural University (CCSHAU), Hisar, Haryana 125004 India

A. K. Dhaka

Karivaradharajan swarnalakshmi.

6 Microbiology, ICAR-Indian Agricultural Research Institute (IARI), New Delhi, 110 012 India

Associated Data

The data that support the findings of this study are available on request from the corresponding author. The data are not publicly available due to private and ethical restrictions.

Delayed sowing of maize hybrids could exacerbate the capability of maximizing the yield potential through poor crop stand, root proliferation, nutrient uptake, and dry matter accumulation coupled with the inadequate partitioning of the assimilates. This study appraised the performance of five recent maize hybrids viz., PMH-1, PJHM-1, AH-4158, AH-4271, and AH-8181 under timely and late sown conditions of the irrigated semi-arid ecologies. Timely sowing had the grain and stover yields advantage of 16–19% and 12–25%, respectively over the late sown maize hybrids. The advanced hybrids AH-4271 and AH-4158 had higher grain yields than the others. During the post-anthesis period, a greater dry matter accumulation and contribution to the grain yield to the tune of 16% and 10.2%, respectively, was observed under timely sown conditions. Furthermore, the nutrient acquisition and use efficiencies also improved under the timely sown. The nutrient and dry matter remobilization varied among the hybrids with AH-4271 and PMH-1 registering greater values. The grain yield stability index (0.85) was highest with AH-4158 apart from the least yield reduction (15.2%) and stress susceptibility index (0.81), while the maximum geometric mean productivity was recorded with the AH-4271 (5.46 Mg ha −1 ). The hybrids AH-4271 and PJHM-1 exhibited improved root morphological traits, such as root length, biomass, root length density, root volume at the V5 stage (20 days after sowing, DAS) and 50% flowering (53 DAS). It is thus evident that the timely sowing and appropriate hybrids based on stress tolerance indices resulted in greater yields and better utilization of resources.

Introduction

Maize ( Zea mays L.), the third most cultivated crop after rice and wheat, can be grown in various soils and climates due to its versatility 1 , 2 . However, delayed sowing after an optimum time can result in reduced yields anomalies, due to aberrant weather events and irregular rainfall. Timely sowing is crucial for maximizing the yield of maize, and growers are concerned about the yield response of maize to sowing dates 3 . In addition, sowing at an optimum time could enhance the profitability of maize by improving the yields, as the crop has extended period to photosynthesize as well as avoid the artificial grain drying at the end of the crop cycle due to various environmental stresses 4 . The increased maize production is the result of improved agronomic management, varietal development 5 , and advances in plant protection measures. Furthermore, yield increment in a particular area is governed by the timely sowing, due to differences in climate and length of the growing season 6 . The supply of assimilate to grain in cereals derived from current assimilation, which supplies directly to kernels, and then remobilises the temporarily stored assimilates in the vegetative plant parts 7 , 8 . Reserve assimilates storage by a stem is strongly influenced by the growing conditions from emergence to the anthesis. Therefore, the grain yield could be buffered by the reserves accumulated in the vegetative portions of the plant pre-anthesis against the unfavourable current assimilation, particularly during the grain filling 8 .

In rice, an extended period of grain filling is the main determinant of grain yield, as it leads to a higher accumulation of dry matter because of greater cumulative mean temperatures and greater solar radiation interception 9 , 10 . The translocation of dry matter stored in the vegetative plant parts pre-anthesis and accumulation of photosynthates post-anthesis determined the grain yield of rice 11 . The dry matter translocated to grains during grain filling from the accumulated dry matter pre-anthesis and accumulation between flowering to physiological maturity acts as a function of grain yield in maize 12 , 13 , and differs among crop species and nutrient inputs 14 . In addition, it has been reported that ~ 85% of total grain dry weight was derived from the photo-assimilation during the grain filling period in maize 11 . Thus, the timely sowing of the genotypes improved the accumulation of photo-assimilates and their remobilization post-anthesis to grains. Further, timely sowing of maize hybrids could results in better interception of the photosynthetically active radiation leading to improved growth and development of the crop.

The identification of genotypes tolerant to stress and non-stress environment have been reported in crops using the indices, such as, stress susceptibility index (SSI), tolerance index (TOL), yield stability index (YSI), and geometric mean productivity (GMP) 15 , 16 . The SSI distinguishes the genotypes showing a minimum reduction in yield under the stress against the non-stress condition 17 , but it fails to identify the genotypes with high yield and stress-tolerant 16 . Further, the TOL indicates that the higher the TOL value more is tolerant to stress resulting in a higher yield potential of the genotypes 18 . Similarly, YSI is yet another index for identifying the stability of genotypes based on the yield under the stress and non-stress environment 19 . The GMP is similar to STI, wherein it indicates that the genotypes with higher GMP could be selected for both the stress and non-stress environment 15 . The above indices for the selection of genotypes have been reported mostly for wheat and other cereals by inducing an environment susceptible to the stress. Therefore, identifying the maize genotypes with higher yield potential under both stress and non-stress (timely and delayed sowing) environments would be a more robust and efficient approach.

In modern high-yielding maize hybrids, nitrogen use efficiency (NUE) has a negative correlation with the grain N concentration (GNC), defined as the grain yield per unit of nitrogen present in soil and applied fertilizer 16 , 20 , 21 . The decline in GNC could be attributed to the leaves staying green, which depends on the enhanced N absorption from the soil post-anthesis and less remobilization from the vegetative plant parts 22 , 23 . While genotypes with greater NUE under varied conditions can be beneficial in terms of protecting the environment, this should be considered when developing new varieties / hybrids and when recommending fertilizers such as N 24 . Moreover, greater N accumulation post-anthesis has a positive correlation with grain yield in rice, indicating that the post-anthesis N-accumulation plays an essential role in expanding the grain yield 25 . Subsequently, adequate concentration of phosphorus (P) is imperative for maintaining a high photosynthesis rate and enhancing the dry matter accumulation 26 . In addition, the extended period of leaf photosynthesis enhanced the grain yield but the trade-off with N and P remobilization in leaves during the grain filling as it shortens leaf photosynthesis 27 . In contrast, potassium (K) differs from P and N as it functions in various enzyme activation, synthesis of protein, maintaining osmotic balance and soluble metabolites translocation within plant tissue 28 . The hidden half of the plant (root systems) greatly influenced the performance of the above-ground portion particularly the formation of grain. Thus, the genotype with better rooting pattern could enhance the acquisition of limited resources (water, nutrients, etc.) and improved the yielding potential of the crop. Further, the root system architecture differs among genotypes, and so too does the nutrients and water uptake and ultimately the yield. Moreover, genotypes with improved root growth parameters and proliferation proportionately partitioned the captured resources and enhanced the yields 29 , 30 .

High-yielding hybrids are a boon for growers, but they require higher inputs as well as timely monitoring of all management practices. Hence, it is necessary to evaluate the performance of advanced/recent hybrids under the optimum nutrient management with varying sowing dates to know the productivity potential. This field study investigates the productivity, translocation and accumulation of dry matter besides the nutrient use-efficiencies and stress tolerance indices under the timely and late sowing of maize hybrids, in irrigated semi-arid ecologies.

Harvest index, grain and stover yields

On an average, the timely sown genotypes recorded 5–10% higher number of cobs per ha over the delayed sowing. While, among the hybrids, advanced hybrid AH-4271 recorded significantly a greater number of cobs ha −1 . Harvest index (HI) did not differ significantly under different sowing times, but advanced hybrids AH-4158 and AH-4271 showed significantly higher HI. Timely sowing gave a significantly stover yield advantage of 12–25.9% over the late sowing. PMH-1 and PJHM-1 had 5.2–16.9% and 5–11% greater stover yields than other hybrids in 2020 and 2021, respectively. With timely sowing of hybrid maize, grain yield increased by 16.7–19.2% compared to late sown hybrids. Hybrid AH-4271 had the highest grain yield, being similar to AH 4158, but significantly greater than the other hybrids (Table ​ (Table1 1 ).

Number of cobs, harvest index and yields of maize genotypes under timely and late sown conditions.

Within sowing time and genotypes, different letters indicate the significant difference ( p  ≤  0.05 ); HI: Harvest index, The S × G implies the interaction between sowing time and genotypes; **Significant at p  ≤  0.01 ; *Significant at P  ≤  0.05 ; ns: not significant.

Yield attributes

The grains cob −1 under the timely sown was higher in 2020, but similar to the late sown in 2021. The highest grains cob −1 was recorded with AH-4271. Further, heavier cobs were harvested in timely sown than in late sown crop, though being similar in 2021. Among hybrids, the PMH-1 recorded the maximum cob weight, being similar to the AH-4158 and AH-4271. The timely sown crop produced 22.7% and 8.2% higher grain weight cob −1 over the late sown crop during 2020 and 2021, respectively. While, the grain weight among the hybrids were comparable, except AH-8181 in 2020, but in 2021 PMH-1 recorded the highest weight over other hybrids (Table ​ (Table2 2 ).

Yield attributes of maize genotypes under timely and late sown conditions.

Within sowing time and genotypes, different letters indicate the significant difference ( p  ≤  0.05 ); HI- Harvest index; The S × G implies the interaction between sowing time and genotypes; **Significant at p  ≤  0.01 ; *Significant at p  ≤  0.05 ; ns- not significant.

Accumulated dry matter, its translocation and contribution to the grain yields

In 2020 and 2021, the late sowing led to a 6.3–14.6% increase in the dry matter translocation before anthesis (Fig.  1 a). Pre DMT translocation was the greatest with AH-8181 and PJHM-1. The effectiveness of dry matter translocation (Pre DMTe) was also higher under late sown than the timely sown crop (Fig.  1 b). While AH-8181 had the greatest Pre DMTe in 2020, PJHM-1 and AH-8181 in 2021. On the other hand, hybrids sown at the right time accumulated 26.5% (2020) and 5.7% (2021) greater dry matter after anthesis (Post DMA) (Fig.  1 c). PMH-1 and AH-4271 recorded the highest Post DMAs. Post anthesis dry matter accumulation efficiency (Post DMAe) under timely sowing was 2.4–19.9% greater than the late sowing (Fig.  1 d). PMH-1 and AH-4271 had a greater Post DMAe than the other hybrids. In 2020, the contribution of pre-anthesis dry matter translocation to grain yield (Pre DMTg) was higher for the late sowing (Fig.  1 e), while in 2021, it was the same for both the sowing dates (Fig.  1 f). In both the years, AH-8181 gave maximum Pre DMTg. Post-anthesis dry matter accumulation contributed 5.6–14.7% more to grain yield under the timely sowing compared to the late sowing. Hybrids AH-4271 and PMH-1 had similar Post DMAg in 2020, while PMH-1 had the maximum in 2021.

Pre DMT- pre-anthesis dry matter translocation ( a ), Pre DMTe- pre-anthesis dry matter translocation efficiency ( b ); Post DMA- post-anthesis dry matter accumulation ( c ), Post DMAe- post-anthesis dry matter accumulated efficiency ( d ); Pre DMTg- pre-anthesis dry matter translocation and Post DMAg- post- anthesis dry matter accumulation contribution to grain yield ( e , f ). Within years, sowing time, and genotypes, different letters on the individual bars of a figure indicate significant difference ( p  ≤ 0.05).

Nutrient concentration in different plant parts at flowering and maturity

Under the timely sown conditions, leaf and stem –N concentration at flowering was 2.9% and 8.8% greater than in the late sown, while hybrids didn’t differ significantly for leaf -N (Table ​ (Table3). 3 ). However, AH-4148 and AH-4271 had greater stem –N. Contrary to the N, the late sown crop had greater leaf and stem -P than timely sown, whereas hybrids had similar stem –P. For leaf –P concentration, PMH-1 being similar to AH-4158 and AH-8181, but greater than PJHM-1 and AH-4271. Leaf -K concentration in timely sown maize was 15.8% greater than that in the late sown. AH-4271 had greater leaf -K than that of PMH-1 and AH-8181, being similar to PJHM-1 and AH-4158. At maturity, the leaf and stem –N under timely sown was 10.7% and 3.8% greater, respectively over the late sown. The late seeded crop had higher leaf –P concentration than the timely sown, and the hybrid AH-8181 accumulated greater leaf –P than the other hybrids. Similarly, the leaf –P concentration was the highest in AH-8181, being comparable to PMH-1 and PJHM-1. While grain P in the hybrids was similar apart from the AH-4158. When sown early, the K concentration (leaf, stem, and grain) in maize hybrids was greater than when sown late. PJHM-1 had the highest leaf and grain –K concentration, while AH-8181 greater stem –K concentration.

Nutrients concentration in different plant parts of maize genotypes at 50% flowering and at maturity under timely and late sown conditions.

N: Nitrogen, P: Phosphorus, K: Potassium. The nutrient concentration given in the table is the mean of 2 years (i.e., 2020 and 2021). Within sowing time and genotypes, different letters indicate the significant difference ( P  ≤ 0.05); the S × G implies the interaction between sowing time and genotypes; **Significant at p  ≤ 0.01; *Significant at p  ≤ 0.05; ns: not significant.

Nitrogen translocation and uptake

Pre-anthesis N translocation in late-sown crop was 2.3–6.4% lower than in the timely-sown crop (Fig.  2 a). In both the years, hybrid AH-4158 exhibited significantly a greater translocation than other hybrids and was comparable to the hybrid AH-8181. The translocation efficiency (Pre NT eff.) was higher with the late-sown to the tune of 8.1–8.6% (Fig.  2 c). The hybrid AH-8181 achieved the best efficiency among the hybrids. After anthesis under the late sown condition, the N uptake (Post Nup) was 6.3% greater; however, timely sown had 12% greater uptake in 2021 (Fig.  2 b). As for N uptake, PJHM-1 was comparable to the AH-4271 and AH-8181, but significantly different from the PMH-1 and AH-4158. In 2021, the uptake of N by AH-4271 was 9.3 and 12.4% greater than that of PMH-1 and AH-8181. The proportion of N uptake after anthesis to total N accumulation (Post NR) varied with years and sowing times (Fig.  2 d). In 2020, PJHM-1 had significantly a greater Post NR than the other hybrids, but in 2021 the hybrids didn’t differ significantly.

Pre NT- pre-anthesis Nitrogen translocation ( a ), Post NuP- post-anthesis Nitrogen uptake ( b ), Pre NT-eff.- pre-anthesis Nitrogen translocation efficiency ( c ), Post NR- ratio of post-anthesis N uptake to total N accumulation ( d ). Within years, sowing time, and genotypes, different letters on the individual bars of a figure indicate significant difference ( p  ≤ 0.05).

Phosphorus (P) translocation and uptake

Pre-anthesis P translocation under the late sown crop was significantly greater than timely sown crop (Fig.  3 a). PMH-1 and AH-4271 had significantly greater pre-anthesis translocation rates than the other hybrids. Under late sowing, the translocation efficiency (Pre PT efficiency) was greater than the timely sowing (Fig.  3 b). In comparison with AH-4158, AH-4271 had significantly a higher Pre PT efficiency. Under timely sown conditions, the P uptake after anthesis (Post Pup) was 8.9–48.9% greater than the late sown crop (Fig.  3 c). In 2020, PJHM-1 and AH-4271 were similar, but significantly greater than other hybrids, whereas in 2021, it was PMH-1 and AH-8181. Under timely sowing, total P accumulation (Post PR) increased by 16.9–49.5% compared to the late sowing (Fig.  3 d). However, among the hybrids, in 2020 Post PR was highest with AH-4271, and in 2021, the PJHM-1 and AH-8181 were comparable, but significantly more than the other hybrids.

Pre PT- pre-anthesis phosphorus translocation ( a ), Post PuP- post-anthesis phosphorus uptake ( b ), Pre PT-eff.- pre-anthesis Phosphorus translocation efficiency ( c ), Post PR- ratio of post-anthesis P uptake to total P accumulation ( d ). Within years, sowing time, and genotypes, different letters on the individual bars of a figure indicate significant difference ( p  ≤ 0.05).

Potassium (K) translocation and uptake

The K translocation (Pre KT), its uptake after anthesis (Post Kup), and the ratio of K uptake to total K accumulation (Post KR) under timely sown were significantly greater than the late sown (Fig.  4 a–d). Compared to the late sowing, timely sown had an increments of 2.9–8.9% (Pre KT), 33.7–43.2% (Post Kup), and 24.5–36% (Post KR), respectively. PJHM-1 had the highest Pre KT and Pre KT efficiency than the other hybrids. Post Kup for AH-4158, AH-4271 and PMH-1 had recorded greater values than other hybrids. In 2020, AH-4158 and AH-4271 had comparable Post KRs, but AH-4271 recorded significantly more than other hybrids, however in 2021, PMH-1 recorded a statistically higher Post KR.

Pre KT- pre-anthesis potassium translocation ( a ), Post KuP- post-anthesis potassium uptake ( b ), Pre KT-eff.- pre-anthesis potassium translocation efficiency ( c ), Post KR- ratio of post-anthesis K uptake to total K accumulation ( d ). Within years, sowing time, and genotypes, different letters on the individual bars of a figure indicate significant difference ( p  ≤ 0.05).

Contribution of nutrient translocation and uptake to the grain yields

Despite of varying sowing dates and years, N translocation before anthesis did not affect the grain yields. Both PMH-1 and AH-4158 showed greater contributions to the pre-anthesis N translocation than did AH-4271 (Fig.  5 a,b). With late sown conditions, pre-anthesis P translocation was 14.8–53.3% greater than the timely sown crop (Fig.  5 c,d). Pre-anthesis translocation was largely accounted by AH-4158, followed by PMH-1. A greater K translocation was recorded under the late-sown conditions (Fig.  5 e,f). Meanwhile, PJHM-1 outperformed the other hybrids; timely sown hybrids contributed 10.8% more N to the grain yield from post-anthesis uptake (Fig.  5 a). In 2021, AH-4271 increased the post-anthesis uptake of N by 9.7–16.3%. Also, timely sowing increased the P uptake by 49.4% and 10.3%, respectively in 2020 and 2021 (Fig.  5 c,d). P uptake contributions were higher for AH-4271 and AH-8181 in 2020, whereas in 2021 they were PJHM-1 and AH-8181. Through post-anthesis K uptake, timely sowing contributed 21.1–29.7% more to grain yield over the late sowing (Fig.  5 e,f). AH-4158, AH-4271 and PMH-1 contributed most to the post-anthesis K translocation compared to other hybrids.

Pre NTg- Pre-anthesis nitrogen translocation and Post NupG- Post-anthesis nitrogen uptake contribution to grain yield ( a , b ); Pre PTg- Pre-anthesis phosphorus translocation and Post PupG- Post-anthesis phosphorus uptake contribution to grain yield ( c , d ); Pre KTg- Pre-anthesis potassium translocation and Post KupG- Post-anthesis potassium uptake contribution to grain yield ( e , f ).

Nutrient uptake in the shoot (above-ground) and efficiencies

Compared to late sown, the timely sown crop had a greater uptake of shoot N by 15.6% (2020) and 29.3% (2021) (Table ​ (Table4). 4 ). The hybrid AH-4271 was most effective when it came to absorbing N in the shoot. PMH-1 and AH-8181 recorded the highest values for shoot P uptake. Further, timely sown hybrids had 21.8% and 32.2% greater shoot K uptake than the late sown hybrids. In both the years, hybrids PMH-1, AH-4271 and PJHM-1 showed a greater shoot K uptake. Timely sown maize had 15.8–29.5% greater N uptake efficiency (NupE) than the late sown maize. In both the years, AH-4271 was the most efficient hybrid in absorbing N. On the other hand, the timely sown crop had a greater P uptake efficiency (PupE) than the late sown crops. The P uptake efficiency (PupE) in hybrids was greater for PJHM-1 in 2020, but for PMH-1 and AH-8181 in 2021. The K uptake efficiency (KupE) under timely sown was 21.7–32% greater than the late sown crop. PMH-1 and PJHM-1 gave the maximum K uptake efficiency in both years.

Above ground nutrient uptake and nutrient uptake efficiencies of maize genotypes under timely and late sown conditions.

AGN: Total above ground nitrogen uptake, AGP: Total above ground phosphorus uptake, AGK: Total above ground potassium uptake, NupE: Nitrogen uptake efficiency, PupE: Phosphorus uptake efficiency, KupE: Potassium uptake efficiency. Within, sowing time and genotypes, different letters indicate the significant difference (p ≤ 0.05); The S × G implies the interaction between sowing time and genotypes; **Significant at ( p  ≤ 0.01); *Significant at p  ≤ 0.05; ns: not significant.

Nutrient (N, P and K) use efficiencies

The N use efficiency (NUE, kg kg −1 ) under timely sown conditions was 16.9–19.9% greater than the late sown conditions (Fig.  6 a). Also, AH-4158 and AH-4271 recorded the greater NUE. Timely sowing had 17.6–19.8% greater P use efficiency (PUE kg kg −1 ). Hybrid AH-4271 had the highest PUE (Fig.  6 b). Under timely sowing, KUE (kg kg −1 ) increased by 17.6–20.2% than the late sown conditions. Again, AH-4271 had the greater KUE and being similar to the AH-4158 (Fig.  6 c).

NUE- nitrogen use efficiency ( a ), PUE- phosphorus use efficiency ( b ), KUE- potassium use ( c ). Within years, sowing time, and genotypes, different letters indicate significant difference ( p  ≤ 0.05).

Stress tolerance indices

In terms of grain and stover yields, maize hybrids exhibited a variable response to the varying stress tolerance indices (Tables ​ (Tables5, 5 , ​ ,6). 6 ). The maximum grain yield reduction of 25.4% was recorded with the hybrid AH-4271. The most stable hybrids for grain yield, however, were AH-4158 and AH-8181. Furthermore, these hybrids also exhibited the least grain stress susceptibility index (SSI). The advanced hybrids AH-4271 (5.52 t ha −1 ) and AH-4158 (5.36 t ha −1 ) had the highest grain mean productivity (MP) and grain geometric mean productivity (GMP), respectively. In addition, the highest stover yield reduction was recorded with the AH-8181 (25.9%) due to the late planting. In terms of stover yield stability index (YSI), AH-4158 and AH-8181 were comparatively more stable with varying sowing times. In contrast, the hybrids with the highest stress susceptibility were PMH-1 and AH-8181. Nevertheless, the MP (7.45 t ha −1 ) and GMP (7.37 t ha −1 ) were highest with the hybrid PMH-1.

Grain stress tolerance indices of maize genotypes under timely and late sown conditions.

*Mean yield of 2 years; YR: Percentage yield reduction, YSI: Yield stability index, SSI- Stress susceptibility index, MP: Mean productivity, GMP: Geometric mean productivity.

Stover stress tolerance indices of maize genotypes under timely and late sown conditions.

*Mean yield of 2 years; %YR: Percentage yield reduction, YSI: Yield stability index, SSI: Stress susceptibility index, MP: Mean productivity, GMP: Geometric mean productivity.

Root system traits

The maize hybrids had a significant variation in the root system traits, such as, root length (RL), root biomass (RB), root length density (RLD), and specific-root length (SRL). PJHM-1 had the highest root length, RLD, and SRL at the 20 DAS, while the maximum root biomass and volume was recorded with the advanced hybrid AH-4271 (Fig.  7 ). At 50% flowering (53 DAS), AH-4271 had the maximum RL, RLD, and SRL (Fig.  8 ), however, PJHM-1 had greater RB compared to other hybrids.

Root morphological traits of five maize hybrids grown in PVC tubes under field condition at V 5 stage (20 DAS). Means followed by different letters on the individual bars/lines of a figure indicate significant difference ( p  ≤ 0.05).

Root morphological traits of five maize hybrids grown in PVC tubes under field condition at 50% flowering (53 DAS). Means followed by different letters on the individual bars/lines of a figure indicate significant difference ( p  ≤ 0.05).

Timely sowing of maize hybrids enhanced the yields through better crop stand, improved yield attributes coupled with higher post-anthesis accumulation of dry matter and nutrient uptake. The contribution of post-anthesis dry matter accumulation to grain yield under both the sowing dates was much higher than the contribution from dry matter translocation. Indeed, the contribution of dry matter translocation to grain yield under the late sown was more than the timely sown, thereby indicates the importance of stored assimilates before anthesis under stress environment. While, for nutrients (N and K), the contribution to grain yield from translocation was more compared to the uptake, but the reverse hold true for P. Dry matter translocation and accumulation, nutrients translocation and uptake, and their contribution to grain yields varied significantly among the hybrids. Further, hybrids with higher geometric mean productivity and tolerance index are more productive under the varying sowing dates. On an average, timely sowing had a 17.9% and 18.9% yield advantage for grain and stover yields, respectively (Table ​ (Table1), 1 ), which could be attributed to the better photosynthates partitioning within the plant, as the crop has an extended period of photosynthesis 4 , with more favourable weather conditions during the growth and development for a particular region 6 . In addition, timely sowing also enhances the synchronization of maximum green leaf area index and the peak solar radiation 31 , thereby, improved the intercepted photosynthesis rate and hence the crop development 32 , resulting in greater yield. Additionally, the higher yield of hybrids in 2021 could be attributed to higher rainfall and its better distribution, particularly during the reproductive stage (Suppl. Figure  1 ).

The delayed sowing had the negative impacts on yields, by reducing the kernel number and their weight 33 . Comparable findings for the reduced kernel number and weight have also been reported by 5 , wherein late sowing would not be able to establish a proper root system under stress conditions. Hence, the uptake and partitioning of water and nutrients under the late sowing couldn’t meet the crop requirement for proper growth and development during the reproductive stage, which might have led to the under developed kernel. Subsequently, delayed sowing could also reduce the number, size and activity of growing grains coupled with the decreased supply of assimilate to grains during the period of grain filling, hence the grain yield 31 . There was a strong correlation between kernel weight with temperature and solar radiation 34 , and kernel weight with the grain yield 31 . Further, the main cause of the reduction in grain yield under late sowing was the reduction in grain number 35 .

Post-anthesis dry matter accumulation under timely sown accounted for 62.5% (av. of 2 yrs.) of the grain yield, wherein it was 55.8% greater than the late sown crop. While, it ranged from 55.6 to 67.7% in 2020 and 48.1 to − 64.1% in 2021 among the hybrids (Fig.  1 e,f), if respiratory losses for maintenance and remobilization of pre-anthesis accumulated assimilate are not taken into consideration 27 . The assimilates for grain formation don’t come entirely from the current assimilation which are directly transferred to the kernels, but also from the remobilization of temporarily stored assimilates in different vegetative plant parts 12 . Accumulated dry matter after anthesis is the major source for grain filling 13 , and in this study, the timely sown accumulated 17.2% higher dry matter and contributed 10.5% greater to the grain yield (Fig.  1 c,d) over the late sown crop. This could in fact be associated with the congenial environment for growth and development 36 , particularly during the grain filling period. In addition, hybrids with greater post-anthesis dry matter accumulation had the positive effects on grain yield, though PMH-1 had relatively lower grain yield which could be ascribed to the lower number of cobs per unit area (Table ​ (Table2 2 ).

The N, P, and K accumulated in vegetative organs of the crop before anthesis, remobilized for grain filling. However, unlike the dry matter, the N, P, and K uptake post-anthesis could not meet the requirement for grain development 27 . Indeed, this study outlined that, the larger amount of N and K uptake occurred before anthesis, while the reverse is true for P (Figs.  2 a,b, ​ a,b,4a,b) 4 a,b) irrespective of the sowing dates and hybrids. The N and P are incorporated into the leaves and along with K it takes part in photosynthesis, hence recycling and remobilization of the stored N, P, and K in the vegetative tissues pre-anthesis would affect the photosynthesis processes 27 , 28 . It has been reported in maize and wheat that a greater amount of N and P accumulated during pre-anthesis is remobilized and recycled under the N and P deficiency 37 , 38 .

In the present study, timely sown had higher post-anthesis uptake of N, P, and K (Figs.  2 b, ​ b,3b, 3 b, ​ b,4b) 4 b) over the late sown crop, which could be the result of better growth and development of both above and the below-ground due to congenial environment, particularly during the grain filling period. Similar results of greater uptake of N, P, and K during the post-anthesis was reported by 39 . Further, the greater uptake of N, P, and K during the post-anthesis implies that it had the priority to be used in grain formation as can be visualized with the greater yields (Table ​ (Table1). 1 ). The greater post-anthesis uptake of N, P, and K under the timely sown crop is used to prolong the stay-green period of leaves, consequently, promotes more grain formation resulting in better yields 22 , 23 , but the grain % N was lower compared to the late sown crop (Table ​ (Table3). 3 ). Similar results of higher post-anthesis nutrient uptake with lower grain % N have also been reported by 20 , 21 , 27 .

The improvement in yields of maize hybrids under variable environmental conditions is desirable, hence in the present study various stress indices were employed to find out the best performing hybrid under the timely and late sown conditions. The higher rate of mean productivity (MP) and geometric mean productivity (GMP) coupled with a lesser stress susceptibility index (SSI) indicated that the genotypes had greater stress tolerance with the enhanced yield potential 15 . The hybrid AH-4158 had the least yield reduction and SSI with greater YSI (Table ​ (Table5), 5 ), thereby, it had greater tolerance with the time of sowing. Subsequently, AH-4271 outperformed for grain MP and GMP though it was more susceptible to late sowing. This hybrid AH-4271 would be well suited under the timely sown with higher productivity potential. In addition, this hybrid had greater stability for the stover yield with different sowing dates (Table ​ (Table6 6 ).

A positive correlation was reported between SSI and grain yield of wheat genotypes to identify the best performing varieties under stress conditions 40 . Further, classifying the genotypes based on MP and GMP were similar, and had positive relations with the grain yield under normal and stress conditions 15 , 41 . The MP, which is the average productivity of yield under stress and normal conditions 42 , and its greater value denotes a better performance of hybrids under the stress conditions, hence a good criterion for selecting a hybrid tolerant to stress. The studies of drought stress on maize hybrids yield reported that under normal and mild stress conditions, the GMP, MP, and stress tolerance index (STI) were important indices for identifying the best performing hybrids under the variable environments 43 . Further, a positive correlation of grain yield with MP and GMP under severe stress and normal conditions were observed, thereby helps in determining the drought-tolerant hybrids 44 .

In our experiment, a hybrid with greater MP, GMP, and YSI could be used for identifying the hybrids adaptability to different sowing dates (timely and late). In terms of grain yield, the AH-4271 and AH-4158 had greater MP and GMP under the variable sowing dates, indicating their superiority. The adoption of N efficient hybrids is a vital management strategy for enhancing the N use efficiency (NUE) 45 . The NUE is grain yield per unit available-N both from soil and through applied fertilizer 16 . In cereals, the NUE has been reported to be about 40% of the applied fertilizers 46 . In our study, the NUE ranged between 10–15 kg kg −1 considering the contribution from 0.0–0.30 m soil profile in addition to the applied N fertilizer (Fig.  6 a).

The greater NUE of AH-4271 and AH-4158 under the timely sown conditions could be attributed to the enhanced uptakes by their root coupled with the better assimilation and remobilization in the shoot 16 , 47 . Studies on wheat showed that NUE could be improved through optimization of the root system 48 , 49 . Furthermore, significantly greater P and K use efficiencies under timely sown (Fig.  6 b,c) might be the result of better growth and development with higher grain yield due to the congenial crop environment. Subsequently, differential growth and development habits of the hybrids in changing environments lead to variation in P and K use efficiencies. The greater P and K efficiencies with the AH-4271 and AH-4158 (Fig.  6 b,c), showed their responsiveness and superiority through better adaptation under the varied ecologies. The importance of growing nutrients use efficient genotypes has been emphasized, as it would reduce the excessive fertilizers input without yield penalty 16 , 49 .

In addition, the rooting traits varied among the hybrids (Figs.  7 , ​ ,8), 8 ), where in the hybrid with better root length and biomass coupled with greater root length density produced better yields (Table ​ (Table1). 1 ). The genotypes with better root proliferation at the early stage might have led to the better crop establishment and used the available resources more efficiently and partitioned proportionately to different plant parts, resulting in the greater output. Indeed, the greater number of cobs per unit area was recorded with those hybrids having better root morphological traits, thereby indirectly implies more crop stand on a unit area. Genotype with a greater root growth and proliferation proportionately partitioned the captured resources and thus enhanced the yields 29 , 30 . Similarly, studies on wheat showed that root biomass was positively correlated to the number of grains spike −1 and yields 50 , so was the case in the present study, i.e., a higher number of grains cob −1 and grain yields related.

Conclusions

Timely sowing, a resource-saving practice plays a vital role in enhancing the yield potential of maize genotypes. Compared with late sown, timely sown yielded 16–19% more grain and produced 5–10% more cobs per hectare. During the grain filling period, post-anthesis dry matter accumulation is crucial, and it was significantly higher with the timely sowing compared to the late sowing. We observed maximum nutrient use efficiency, nutrient uptake, and nutrient contribution to grain yield under the timely sowing conditions. Advanced hybrids, AH-4271 and AH-4158, performed better in various parameters, viz., grain yield, nutrient uptake, and cobs per ha. Hybrid AH-4158 showed higher yield stability, a lower stress susceptibility index, and a lower percentage of grain yield reduction, indicating the enhanced capacity for flexibility in a variety of crop-growing conditions, together with hybrid AH-4271. Indeed, better root morphology correlates with the greater nutrient use efficiency, dry matter accumulation, and nutrient remobilization to produce higher yields. Therefore, assessing hybrids based on stress indices, nutrient remobilization, and grain yield could lead to identifying the best hybrids under the variable crop conditions.

Materials and methods

Experimental site and weather conditions.

A Fixed-site field experiment was conducted for 2 years at the ICAR-Indian Agricultural Research Institute, New Delhi, India, planting maize genotypes under the timely and late sown conditions during the rainy seasons of 2020 and 2021. The region falls under the Trans Indo-Gangetic plains with 28°38′ N latitude, 77°10′ E longitude, and 229 m amsl. The climate is semi-arid with hot summers and rainy in monsoon (July–September), with scattered rains in winter. There is an annual mean rainfall of 650 mm, and the mean maximum and minimum temperatures range from 20–40 °C to 4–28 °C, respectively. The 2 years weather observations (2020–21) recorded by the automated observatory in the adjacent experimental site are summarized in suppl. Figure  9 . Before preparatory tillage, the soil samples were collected randomly from 0.0–0.15 m to 0.15–0.30 m undisturbed soil depth. the samples were air-dried, ground, sieved through a 0.2 mm sieve, and stored in air-tight polyethene bags for further analysis of soil chemical properties, viz. soil pH (1:2.5, soil: water 51 ), KMnO 4 -oxidizable N 52 , NaHCO 3 extractable P 53 , NH 4 OAc exchangeable K 54 , and soil organic carbon (S OC ) 55 (Table ​ (Table7 7 ).

Initial soil chemical properties of the experimental site.

Cultural operations, experimental design, and crop management

Pre-sowing irrigation was applied before preparatory tillage operations. The field was deep ploughed twice using a disc harrow (0.00–0.20 m depth) followed by planking with a rotavator/ cultivator twice and finally levelled. The experiment was laid out in a split-plot design with three replicates. Two sowing times (timely and late sown) were allocated to the main plots and five recently released maize hybrids (PMH-1, PJHM-1, AH-4158, AH-4271, and AH-8181) to sub-plots, with a sub-plot size of 20 m 2 (4 m × 5 m). In 2020, the crop was sown on 6th July (timely) and 27th July (late), while in 2021 it was sown on 3rd July (timely) and 24th July (late), respectively. The seeds were dibbled manually at a spacing of 0.75 m (row–row) × 0.20 m (plant–plant) in both the seasons. Earthing-up was done at the knee-high stage for better crop growth, prevent lodging and uniform distribution of irrigation water. Based on the critical growth stages and the rainfall received during the crop seasons, irrigation water was applied to a depth of 0.05 m. The recommended fertilizer application rate for maize was 150:26.2:49.59 kg NPK ha −1 . Nitrogen (N) was applied as urea (46% N), phosphorus (P) and potassium (K) through di-ammonium phosphate (46% P 2 O 5 ), and muriate of potash (60% K 2 O), respectively. At sowing, full doses of P, K, and 50% N were applied uniformly in all the plots, while the remaining 50% N was top-dressed in two equal splits at knee-high and tasseling stages. Weeds were controlled through broad-spectrum pre-emergence herbicide atrazine (50% WP, at 750 g a.i. ha −1 ) applied a day after sowing, followed by one hand weeding at 35–40 days after sowing (DAS). For insect-pests management, particularly fall armyworm ( Spodoptera frugiperda ), a systemic insecticide emamectin benzoate (50% SG, at 200 g ha −1 ) was first sprayed at 20 DAS, followed by need-based at 15–20 d intervals on their appearance. The insect-pests and disease management were carried out uniformly in all the plots based on the recommended practices.

Plant sampling and their laboratory analysis

Plant samples were collected at two stages of the crop growth (tasseling and maturity) at two different dates in each season. In each plot, three plants were randomly cut at the base and separated into leaf, stalks (stem + leaf sheaths + tassel), and leaf, stalks (stem + leaf sheaths + tassel + husk), cob and grain at maturity. The samples were placed in a perforated brown paper bag, air-dried for 48 h, and then oven-dried at 65 ± 2 °C to a constant weight. The dry matter (DM) values were used to determine the translocation, accumulation, and efficiency as per the equations described by (1) 11 , 21 , 56 . A Macro Wiley-mill having a 40-mesh sieve was used for grinding the plant samples, and appropriate amounts (0.5 g) of the ground samples were used to determine the total N concentration employing the modified Kjeldahl digestion process, total P by colored Vanado-molybdo-phosphoric acid procedure, and total K by flame photometer method 57 . The nutrient translocation, uptake, and efficiency were computed in accordance with the Eq. (2) 21 , 58 . The soil available nutrients in this study were determined to the depths of 0.00–0.30 m. Also, the above-ground nutrient uptake at maturity, uptake efficiencies were computed by using Eq. (3) 59 , 60 . The above-ground nutrient uptake was worked out by multiplying the nutrient concentration in stalks and grains with the biomass yields. To estimate the uptake and use efficiencies, the soil nutrients available to a depth of 0.30 m (0.00–0.15 m, 0.15–0.30 m) were used.

Pre NT, Pre NTe, Post Nup, Post NR, Pre NTg, Post NupG- Nitrogen; Pre PT, Pre PTe, Post Pup, Post PR Pre PTg, Post PupG- Phosphorus; Pre KT, Pre KTe, Post Kup, Post KR, Pre KTg, Post KupG- Potassium.

AGN, NupE, NUE—Nitrogen; AGP, PupE, PupE—Phosphorus; AGK, KupE, KUE—Potassium.

The maize genotypes were subjected to mathematical relationships on the basis of stover and grain yields for identifying the best performing genotype under normal and the delayed sowing conditions. The following formulae were used to find out the efficient genotypes 15 , 16 .

• i. Percentage reduction of yield (YR; %) = (Y T  − Y L )/(Y T  × 100)
• ii. Mean productivity (MP) = (Y T  + T L )/2
• iii. Stress susceptible index (SSI) = (1 − Y T /Y L )/SI. Where Stress intensity (SI) was calculated as, SI = 1 − (X T /X L )
• iv. Geometric mean productivity (GMP) = √(Y T  × Y L )
• v. Yield Stability Index (YSI) = Y T /Y L

Where, Y T and Y L are the yields of genotypes under timely and late sown, respectively. The X T and X L denote mean yield of all genotypes under timely and late sown conditions, respectively.

Yield attributes and yields

The physiological growth stages were marked when 50% of the plants attained their particular stage, such as, tasseling, silking, and physiological maturity. In the first season, maize was harvested on 18th and 27th October 2020, while in the second season on 16th and 27th October 2021, respectively for the timely and late sown conditions. The crop was harvested from the middle three rows (4 m × 2 m, 8 m 2 ) leaving two border rows on each side. First, the cobs were hand-picked, then the stover was cut from the ground surface. The harvested produce was sun-dried for 25–30 d to bring down the grain moisture from 20–22% to 14–15% for threshing and the yield measurement for grain and stover were done separately. Further, the yield attributes, such as, number of rows cob −1 , number of grains cob −1 , cob weight (g), grains weight cob −1 (g), and 100-grains weight (TW, g) were determined from the five randomly picked cobs.

The plants were grown in 25 mm PVC tube, 0.195 m diameter, 0.5 m and 1.0 m deep, transparent cylindrical polyethylene sheet placed on the inner PVC tube, filled with soil to a depth of 1.0 m. The PVC tubes were placed vertically on the steel stands in an open field condition. The soil (0.0–0.15 m depth) for the above purpose was collected from the experimental field of the current study. The 5 mm sieve was used to sieve out the debris and other undesirable materials before being packed into the PVC tubes. The soil in the tubes was manually watered carefully to saturation level so as to avoid the excess drainage (Pooniya et al. 30 ). The soil in the tubes was fertilized at sowing to the depth of 0.01 m with an equivalent to the recommended fertilizer rate for maize hybrids (150:26.2:49.59 kg NPK ha −1 ), in the form of urea, diammonium phosphate, and muriate of potash. The plants were hand watered based on the critical growth stages as per the requirement. Three seeds each of the five recent/advanced hybrids were manually dribbled to a depth of 0.05 m in each tube, during the first week of July 2019. Five days after germination, a single plant was maintained in each tube after carefully thinning out the remaining seedlings. The five hybrids were grown in a completely randomized block design with three replicates and harvested twice {V5 stage-20 DAS (Fig.  9 a) and 50% flowering- 53 DAS (Fig.  9 b)}. At first sampling (0.5 m depth tube), the shoots were first cut from the roots at the crown, while in the second (1.0 m depth tube), the shoots and stilt roots above the soil were cut before washing. The polyethylene sheet was gently pulled from the tubes, cut open and dipped in the still water for an hour, repeatedly washed the soil on a 2 mm sieve to produce a clean root sample 29 . The recovered roots were placed in the plastic bags and stored at 5 °C until the scanning of its morphological traits. The scanned images of the roots were analyzed with WinRHIZO professional software (LA2400, Regent instrument, Quebec, Canada) for recording the root morphological traits (total root length and root volume, in the present study). The scanned roots were further dried in a hot air oven at 65 ± 5 °C until obtaining the constant weight for root dry biomass. The root length density was computed by dividing the total root length with the soil volume (0.0149 m 3 —1st sampling and 0.0298 m 3 —2nd sampling). While the specific root length was calculated as the total root length divided by the root biomass 30 .

Destructively measured root growth pattern of the maize genotypes (PMH-1, PJHM-1, AH-8181, AH-4271, AH-4158) grown in PVC tubes under the field conditions at V5 stage (20 days after sowing, DAS) ( a ) and at 53 DAS ( b ).

Statistical analysis

With SAS 9.4, the data were statistically analyzed for split-plot designs through analysis of variance (ANOVA) 61 . In addition, Tukey’s honestly significant difference at 0.05 probability ( p  ≤ 0.05) was used to compare the mean effects of the treatments i.e., sowing dates and genotypes.

Authors have confirmed that all the plant studies were carried out in accordance with relevant national, international or institutional guidelines.

Supplementary Information

Acknowledgements.

We acknowledge to the ICAR-Indian Agricultural Research Institute and the Indian Council of Agricultural Research, New Delhi for providing the necessary field and laboratory facilities during the experimentation.

Author contributions

R.R.Z., N.B., V.P., led the research work, planned, supervised, and conducted field experiments, and read and edited the manuscript. N.B., R.R.Z., K.K. L., A.B., collected soil/plant samples and performed chemical analysis, also wrote the initial draft of the manuscript, and prepared figures, and tables. D.K. Y.S., A.D., G.M., R.K.P., P.G., S.B., A.B., K.S., project supervision, reviewed, read and edited the manuscript with significant contributions. R.D.J. and A.K.D. performed the statistical analysis and prepared figures.

Data availability

Competing interests.

The authors declare no competing interests.

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The online version contains supplementary material available at 10.1038/s41598-023-28224-9.