Does water availability affect the critical N dilution curves in crops? A case study for maize, wheat, and tall fescue crops

doi:10.1016/j.fcr.2021.108301

Field Crops Research

Volume 273, 1 November 2021, 108301

https://doi.org/10.1016/j.fcr.2021.108301 Get rights and content

Highlights

•
Re-analysis of published field crop data shows no conclusive evidence of water availability on the nitrogen dilution curve.
•
The review of the literature provides information that WSC accumulation may be an indirect consequence of a N deficiency.
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Water availability presents limited effect on the allometry of plant mass allocation.

Abstract

Previous studies reported an effect of water deficit on crop nitrogen (N) demand but the statistical significance of this effect is unclear and the physiological mechanism explaining this effect remains poorly understood. The specific aims of this review are (i) assess whether the parameters of the N dilution curve are significantly influenced by the level of water availability and (ii) provide a synthesis of the plausible physiological mechanisms involved on changes in N dilution. To address the first goal, several datasets assessing the effects of water availability on N status of maize (Zea mays L.), wheat (Triticum aestivum L.), and tall fescue (Lolium arundinaceum (Schreb.) Darbysh.) were combined and then used to fit N dilution curves with a Bayesian statistical approach. Concerning the second objective, we investigate two possible mechanisms, namely a modification in water soluble carbohydrates (WSCs) and an effect on plant architecture and morphology impacting the allometry between structural and metabolic compartment. Our re-analysis of published data for tall fescue, wheat, and maize reveals no conclusive evidence supporting an effect of water availability effect on the N dilution curve. In addition, we show that WSCs accumulation under water deficit condition is unlikely to be accompanied by N-rich soluble compounds, and that a reduction of water availability has limited effect on the allometry of plant mass allocation. Lastly, our conclusion provides new insights on the use of a common critical N dilution curve (one model across water levels) for estimating the crop N status where both water (under no extreme deficiency) and N stresses are operating in order to separate direct effect of water availability from its indirect effect on crop N nutrition status.

Introduction

Plant nitrogen (N) status can be robustly diagnosed by the calculation of the N nutrition index (NNI), expressed as the ratio of the actual plant N concentration (%N) and the critical %N (%Nc) at a similar aerial biomass, W (e.g., Plénet and Lemaire, 2000; Gastal et al., 2014; Lemaire and Ciampitti, 2020). The NNI is based on the critical N dilution curve, portraying temporal decreases of plant N demand, i.e. N uptake per unit of crop mass, as plant aged (Lemaire and Salette, 1984). The %Nc defines the minimum %N to maximize plant growth, mathematically expressed with crop W and two parameters (herein termed as A₁ and A₂), and predicted from the critical N dilution curve [%Nc = A₁ W ^−A₂, theoretical framework developed by Greenwood et al. (1990); Lemaire and Gastal (1997), and Lemaire et al. (2008)]. This critical N dilution model has been studied in many crops for segregating NNI < 1 versus NNI> = 1 scenarios (e.g. Justes et al., 1994; Colnenne et al., 1998; Plénet and Lemaire, 2000; Bélanger et al., 2001; Ata-Ul-Karim et al., 2013).

The critical N dilution curve can be interpreted as the result of two processes: i) an allometry between the metabolic plant biomass fraction W_m (the mass of plant tissues directly involved in plant growth process: photosynthetic parenchyma and meristematic tissues) having a high %N (%N_m), and the structural fraction W_s (the mass of plant tissues involved in plant architecture: fibers and vascular bundles) having a low %N (%N_s), as proposed by Caloin and Yu (1984) and Charles-Edwards et al. (1987). As plants get older, the ratio W_m/W_s declines allometrically with W leading then to a decline of %N; and ii) a non-uniform distribution of N within dense canopies, with preferential allocation of N to well illuminated leaf areas at the top of the canopy (Field, 1983; Hirose and Werger, 1987; Lemaire et al., 1991) and N recycling from bottom to top leaf layers that leads to a progressive decline in %N_m as canopy develops.

Water availability effects on critical plant %N and N dilution in crops has been suspected after the result reported in Errecart et al. (2014) on tall fescue of a reduction of %Nc due to soil water deficit. Two hypothesis could explain this reduction of plant N demand, i.e.; the minimum crop N uptake for achieving maximum crop mass: (i) a physiological adaptation of plant linked to osmotic adjustment and an accumulation of free N compounds in plants in water deficit conditions leading then to a more rapid %N decline as proposed by Hoogmoed and Sadras (2016); and (ii) a morphological adaptation of plants through a change in the W_m/W_s allometry in response to water stress. Thus, it is still uncertain if the effect of water deficit reduces critical %N for given value of W.

In this review, we aimed to (i) assess whether the parameters of the N dilution curve are significantly influenced by water availability and (ii) provide a synthesis of the plausible physiological mechanisms involved on changes in N dilution. To address the first goal, several datasets assessing the effects of water availability on N status of wheat (Triticum aestivum L.), maize (Zea mays L.), and tall fescue (Lolium arundinaceum (Schreb.) Darbysh.) were combined and then used to fit N dilution curves with a Bayesian statistical approach. Concerning the second question, we investigate two possible mechanisms, namely a modification in water soluble carbohydrates (WSCs) and an effect on plant architecture and morphology impacting the allometry between structural and metabolic compartment.

Section snippets

Data acquisition

We conducted a literature search using the Web of Science database for studies reporting information on both leaf area index (LAI) and W and %N and W, under contrasting water availability (irrigated vs. rainfed). There were no restrictions in the year of publication or country. The structure of the search to study LAI-W allometry in maize included the terms “nitrogen” or “N”, “maize” or “corn”, "drought", "water stress" or "deficit irrigation", and “leaf area”. To study %N–W allometry, the

Is there an effect of water availability on the dilution curve when the uncertainty of the estimated parameters of the fitted curves is taken into account?

In order to provide a formal answer to this question, a dataset including %N–W observations under rainfed and irrigated conditions for maize, wheat, and tall fescue was analyzed using a Bayesian statistical method (Makowski et al., 2020; Ciampitti et al., 2021). We used a Bayesian statistical method here because our main objective is to analyze the uncertainty in the estimated values of the dilution curve parameters. The Bayesian approach is recognized as being very well adapted to uncertainty

Are the increases in WSC due to reductions of water availability inducing a corresponding decrease in critical %N in plant?

In support of a negative effect of water limitation on plant critical %N, Hoogmoed and Sadras (2016) provided a modified theoretical framework to account for an effect of WSC accumulation in N dilution process. They proposed to add a third plant compartment, the biomass associated to WSC accumulation (Wwsc) to the two other plant compartments: metabolic (W_m) and structural (W_s) of the Caloin and Yu (1984) model. As by definition %N of Wwcs is 0, any variation in size of this component must then

Are architectural and morphological modifications of plant as induced by water deficit susceptible to determine changes in critical %N dilution curves?

According to Lemaire et al. (2007), the dry mass allocation pattern between W_m and W_s of plants as plant size increases is driving the N dilution. For example, Lemaire et al. (1989) and Lemaire and Bélanger (2020) showed that the leaf/stem W ratio of alfalfa (cultivated as a vegetative plant and harvested before flowering) declined allometrically with the accumulation of aboveground crop mass, and that water deficit does not modify the scaling coefficient of this decline despite its large

Limitations and future prospects

This review and the main implications are limited by the availability of data reporting %N, W, and other relevant plant and environmental traits (e.g., LAI, WSC, ET, water supply). In addition, many studies do not report source of error or variation for those critical determinations, limiting the use of the data. Moreover, studies investigating N dilution in plants should provide a clear indication that the value of Wmax was achieved under a non-limiting N nutrition status (testing sufficient

Conclusions

This review provides new insights for a more rigorous assessment of the uncertainty of estimates of the N dilution curves under different water availability, providing examples here for tall fescue, maize, and wheat field crops. From a perspective of changes in WSC and its effect on critical plant %N dilution, the review of the literature provides information that WSC accumulation may be an indirect consequence of a N deficiency rather than a cause of the decline in plant %N. In addition, the

CRediT authorship contribution statement

Ignacio A. Ciampitti: Conceptualization, Methodology, Formal analysis, Investigation, Writing - original draft, Visualization, Supervision, Project administration, Funding acquisition. David Makowski: Formal analysis, Methodology, Writing - review & editing. Javier Fernandez: Data curation, Writing - review & editing, Formal analysis, Visualization. Josefina Lacasa: Data curation, Writing - review & editing, Formal analysis, Visualization. Gilles Lemaire: Conceptualization, Methodology, Formal

Declaration of Competing Interest

The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.

Acknowledgments

The authors gratefully acknowledge the financial support provided by Fulbright Program, the Argentine Ministry of Education for sponsoring J.A. Fernandez’s studies and Kansas Corn Commission for supporting J. Lacasa’s studies and Dr. I.A. Ciampitti’s research program, and the Cland Convergence Institute (16-CONV-0003). This is contribution no. 22-084 from the Kansas Agricultural Experiment Station.

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Citation Excerpt :
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