Environmental and Experimental Botany 75 (2012) 258–267
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Environmental and Experimental Botany
j o ur nal homep age : www.elsevier.com/locate/envexpbot
Restricted root-to-shoot translocation and decreased sink size are responsible for
limited nitrogen uptake in three grass species under water deficit
V. Gonzalez-Dugo
c,1,2
, J.L. Durand
a,∗,2
, F. Gastal
a,2
, T. Bariac
b,2
, J. Poincheval
a,2
a
Unité de Recherche Pluridisciplinaire Prairies et Plantes Fourragères, INRA, BP6, F-86600 Lusignan, France
b
UMR 7618 BIOEMCO, Université Pierre-et-Marie-Curie–CNRS–INRA, Centre de recherche INRA–INA PG, Bâtiment EGER, 78850 Thiverval-Grignon, France
c
Instituto de Agricultura Sostenible – CSIC, Alameda del Obispo, s/n 14004, Córdoba, Spain
a r t i c l e i n f o
Article history:
Received 8 July 2010
Received in revised form 11 July 2011
Accepted 13 July 2011
Keywords:
Dactylis glomerata
Festuca arundinacea
Growth
Lolium multiflorum
Nitrogen concentration
Split-root
a b s t r a c t
Earlier studies showed that water deficit reduces nitrogen (N) uptake and N nutrition index of grasses.
So far, the main effect of water deficit on N uptake and N nutrition status was ascribed to the alteration
of the transpiration-dependent transport of mineral N in the soil solution. A split-root experiment was
performed to determine whether plant and/or solution water potential could alter N uptake and allo-
cation, independently of N fluxes in the soil solution. The split-root experiment allowed to manipulate
separately the water and the N status of the root environment and of the plant, by various combina-
tions of addition of polyethylene glycol 6000 (PEG) and mineral N on half or on the entire root system.
Tall fescue, Italian ryegrass and cocksfoot, known for their contrasted sensitivities to water deficit, were
studied. The addition of PEG largely reduced water and N uptake of the roots on which it was applied
(half or entire root system). A significant accumulation of N was observed on the roots to which PEG
was added, particularly in cocksfoot and tall fescue, hypothetically contributing to the alteration of N
uptake. Cocksfoot displayed a high plasticity in N allocation to leaves related to sheaths, allowing the
maintenance of N concentration and therefore its N status despite the reduction in N uptake. By contrast,
leaf N concentration and N status of tall fescue and Italian ryegrass was more sensitive to water deficit,
similarly to observations of the effect of drought in field studies. Therefore, it is concluded that the effect
of soil solution potential on N uptake and plant N allocation may also contribute to the observed effect
of drought on N status of grass crops, additionally to the effect of drought on the transfer of mineral N in
the soil solution to the root surface. However, the importance of this physiological effect varies among
species.
© 2011 Elsevier B.V. All rights reserved.
1. Introduction
It has repeatedly been observed that soil water deficit alters crop
nitrogen (N) nutrition, even if mineral N is fully available in the soil,
leading to a lower shoot N concentration, a lower crop N status,
and therefore to a crop N deficiency (Lemaire and Denoix, 1987;
Onillon et al., 1995; Gonzalez-Dugo et al., 2005, 2010). Once soil
water deficit is removed, N uptake is restored if N is still available
in the soil (Buljovcic and Engels, 2001; Gonzalez-Dugo et al., 2005).
Crop N uptake is regulated by N availability at the root surface,
which depends on soil water content and on soil water movement
through both mass flow and diffusive water fluxes (Gonzalez-Dugo
∗
Corresponding author. Tel.: +33 5 49 55 60 94; fax: +33 5 49 55 60 68.
E-mail address: jldurand@lusignan.inra.fr (J.L. Durand).
1
Present address.
2
With the technical collaboration of G. Bardou
b
, P. Biron
b
, L. Cousson
a
,
A. Eprinchard
a
, G. Millet
a
, P. Richard
b
, J.-P. Terrasson
a
(see Appendix A).
et al., 2010; Durand et al., 2010). Garwood and Williams (1967)
demonstrated the interaction between local water and N content
of soil layers and the growth of perennial ryegrass in the field, by
injecting nutrients at controlled depths in soil. During soil water
deficit, the decline in soil N supply to roots may induce a crop N
deficiency that contributes, together with the direct effect of plant
water deficit per se, to the overall resulting effect of soil water
deficit on crop growth (Gonzalez-Dugo et al., 2010).
However, in addition to the effect of the water-dependent trans-
port of mineral soil N to the root surface, water deficit also has
major physiological effects on N uptake and N economy of the
plant. By experimentally manipulating root water uptake and plant
water fluxes independently of N availability at root surface (PEG,
split-root experiments), several studies showed that a reduction
in plant water uptake may limit N uptake (Nicolas et al., 1985;
Larsson, 1992; Buljovcic and Engels, 2001; Cayet, 2001). Further-
more, beyond affecting N uptake, water deficit may also alter N
assimilation and N allocation between plant organs. Water deficit
alters nitrate reductase activity (Larsson, 1992, Azedo-Silva et al.,
0098-8472/$ – see front matter © 2011 Elsevier B.V. All rights reserved.
doi:10.1016/j.envexpbot.2011.07.009