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Responses to Drought and Phosphorus Deficiency-And PHYTOHORMONE

Responses to Drought and Phosphorus Deficiency-And PHYTOHORMONE

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2005 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim 1436-8730/05/0408-531
J. Plant Nutr. Soil Sci.
2005,
168,
531–540
DOI:
10.1002/jpln.200520507 531
Plant responsesto drought and phosphorus deficiency:contribution ofphytohormones in root-relatedprocesses
Lutz Wittenmayer
1
*
and
Wolfgang Merbach
11
Martin-Luther-UniversitätHalle-Wittenberg, LandwirtschaftlicheFakultät, Institut für Bodenkundeund Pflanzenernährung,Adam-Kuckhoff-Straße 17b, D-06108 Halle (Saale), GermanyAccepted June 11, 2005
PNSS P05/07P
Summary—Zusammenfassung
Environmental stresses are one of the most limiting factors inagricultural productivity. A large portion of the annual cropyield is lost to pathogens (biotic stress) or the detrimentaleffects of abiotic-stress conditions. There are numerousreports about chemical characterization of quantitatively sig-nificant substrate fluxes in plant responses to stress factorsin the root-rhizosphere system,
e.g.
, nutrient mobilization,heavy-metal and aluminum immobilization, or establishmentof plant-growth-promotingrhizobacteria(PGPR) by exudationof organic anions, phytosiderophores, or carbohydrates intothe soil, respectively. The hormonal regulation of theseresponses is not well understood. This paper highlights thiscomplex process, stressing the involvement of phytohor-mones in plant responses to drought and phosphorus defi-ciency as examples. Beside ethylene, abscisic acid (ABA)plays an important role in drought-stress adaptation of plants.This hormone causes morphological and chemical changesin plants, ensuring plant survival under water-limited condi-tions. For example, ABA induces stomata closure, reductionin leaf surface, and increase in root : shoot ratio and, thus,reduction in transpiration and increase in soil volume forwater uptake. Furthermore, it supports water uptake in soilwith decreasing water potential by osmotic adjustment. Suit-ability of hormonal parameters in the selection for improvingstress resistance is discussed. Auxins, ethylene, and cytoki-nins are involved in morphological adaption processes tophosphorus (P) deficiency (increase in root surface,
e.g.
, bythe formation of more dense root hairs or cluster roots).Furthermore, indole-3-acetic acid increases root exudationfor direct and indirect phosphorus mobilization in soil. Never-theless, the direct use of the trait “hormone content” of a par-ticular plant organ or tissue, for example the use of thedrought-stress-induced ABA content of detached leaves inplant breeding for drought-stress-resistant crops, seems tobe questionable, because this procedure does not considerthe systemic principle of hormonal regulation in plants.
Keywords:
phytohormones/droughtstress/phosphorusdeficiency/ rhizosphere / root
Reaktionen von Pflanzen auf TrockenstressundPhosphormangel: Die Rolle von Phytohormonen inwurzelbezogenen Prozessen
Umweltstress stellt den wesentlichsten Limitierungsfaktor fürdie landwirtschaftliche Produktion dar. Ein erheblicher Teilder jährlichenErnten geht durch pathogeneOrganismen (bio-tischer Stress) oder durch die verheerende Wirkung abioti-scher Stressoren verloren (v. a. Trockenstress und Nährstoff-mangel). Es gibt zahlreiche Untersuchungen zur stofflichenCharakterisierung der pflanzlichen Stressreaktion an derWurzel, z.B. Nährstoffmobilisierung, Schadstoffimmobilisie-rung oder Etablierung von wachstumsfördernden Rhizobak-terien durch Wurzelabscheidungen. Die hormonelle Steue-rung dieser Prozesse ist bisher weniger erforscht. Der Artikelgeht dieser Problematik am Beispiel von Trockenstress undPhosphormangel unter besonderer Berücksichtigung vonPhytohormonen nach. Bei der Anpassung von Pflanzen anWassermangelbedingungen spielt neben Ethylen das Phyto-hormon Abscisinsäure(ABA) eine wichtigeRolle. Es induziertmorphologische und chemische Veränderungen in derPflanze, die ein Überleben unter Wassermangelbedingungenermöglichen. Beispielsweise induziert die ABA den Stomata-schluss, eine Verringerung der Blattoberfläche sowie eineErhöhung des Wurzel:Spross-Verhältnisses und bewirktdadurch eine verringerte Transpiration und Vergrößerung desBodenvolumens zur Erschließung von Wasservorräten. Da-rüber hinaus kann eine ABA-induzierte Anreicherung vonosmotisch wirksamen Verbindungen zur Wasseraufnahmebei sinkendem Wasserpotential im Boden beitragen. BeiPhosphat (P)-Mangel sind vor allem Auxine, Cytokine undEthylen an der morphologischen Anpassung der Wurzeln(Vergrößerung der Wurzeloberfläche durch verstärkte Bil-dung von Wurzelhaaren oder Proteoidwurzeln) beteiligt. Da-rüber hinaus bewirkt Indolyl-3-Essigäure eine Intensivierungder Abgabe von Wurzelabscheidungenzur direkten oder indi-rekten P-Mobilisierung in der Rhizosphäre. Trotzdem wird dieunmittelbare Verwendung des Indikators „Hormongehalt“eines bestimmten Pflanzenorganes, beispielsweise dertrockenstressinduzierte ABA-Gehalt von abgeschnittenenBlättern, für die Züchtung auf Stressresistenz als problema-tisch angesehen, da sie das systemische Prinzip der Hor-monregulationnicht berücksichtigt.
1 Introduction
Knowledge about plant responses to various stress factorsand the relationship between suboptimal environmental con-ditions and plant growth and crop yield is very important forseveral reasons: First, land areas affected by various stressfactors have to be used in agricultural production at present.Furthermore, regions with partially very low suitability for agri-cultural use (
e.g.
, marginal soils, zones with water limitation)are the only way for expansion of field areas and, thus, forincreasing food production necessary for a growing worldpopulation (
European Parliament 
, 1999;
Smithson 
and
Sanchez 
, 2001;
Williams 
, 2002). Second, climatic changes(
e.g.
, long-term increase in temperature with simultaneousdecrease in precipitation in central and eastern Germany)
* Correspondence: Dr. L. Wittenmayer;e-mail lutz.wittenmayer@landw.uni-halle.de
 
may result in more frequent stress events even in regionstraditionally used for agricultural purposes (
BMBF 
, 2003;
Gerstengarbe 
et al., 2003;
Hulme 
and
Sheard 
, 1999;
Umwelt- bundesamt 
, 2002).In Africa, South Asia, and Australia, half of the total landareas are affected by water deficiency, while in South Amer-ica and Southeast Asia, mineral imbalances are the dominat-ing factors. Thus, drought and chemical constrains are themajor limitations to agricultural use of land (
Smithson 
and
Sanchez 
, 2001). After nitrogen (N), P is usually the most lim-iting nutrient for crop production (
Schachtman 
et al., 1998).Therefore, due to their importance in agricultural production,drought and P deficiency are illustrated here in more detail.Plants develop a wide range of strategies to cope with stresssituations. Under conditions of water deficiency, droughtescape and drought tolerance are two important strategies toensure plant growth (
Bänziger 
et al., 2000). In the first case,plants finish their growth period before drought stress mayaffect yield formation. In plant breeding, this strategy is con-sidered by selection for earliness. Drought tolerance may beincreased by morphological or chemical changes of plants,
e.g.
, stomatal regulation (
Wilkinson 
and
Davies 
, 2002),reduction of the plant surface, increased root : shoot ratio,accumulation of osmolytes (
Chen 
and
Murata 
, 2002;
Nayyar 
and
Walia 
, 2004), or rhizosphere processes (
McCully 
and
Boyer 
, 1997). For example,
Walker 
et al. (2003) speculatedthat root exudates could play a major role in the maintenanceof root-soil contact, which is especially important to the plantunder drought and drying conditions, when hydraulic continu-ity will be lost.
Young 
(1995) found that rhizosheath soil wassignificantly wetter than bulk soil and suggested that exu-dates within the rhizosheath increase the waterholding capa-city of the soil.Under P-stress conditions, strategies for improving phos-phorus-use efficiency are: first, increasing the root surface-soil contact area by modifying root morphology; second,increasing the effective root area by root symbiosis witharbuscular mycorrhizal fungi; and third, increasing nutrientavailability through rhizosphere modification (
e.g.
, shiftingpH, release of reductans or chelators) (
Hinsinger 
et al., 2003;
Jones 
et al., 2004;
Randall 
et al., 2001;
Rubio 
et al., 2003;
Ryan 
et al., 2001;
Schilling 
et al., 1998;
Vance 
et al., 2003).The use of these traits in plant breeding, particularly the im-provement of root morphology, is time-consuming and labor-intensive and, therefore, limits its application in breeding pro-grams where large numbers of genotypes need to bescreened (
Ortiz-Monasterio 
et al., 2001;
Sinclair 
and
Vadez 
,2002). Due to their better accessibility, shoot-related para-meters were mostly considered in the selection processes inpast decades. There is much less information about rootcharacteristics important for stress resistance. Particularlythe regulation of stress responses in the root-rhizospheresystem is not well understood. Doubtless, phytohormonesplay an essential role in these processes (
Casson 
and
Lindsey 
, 2003;
Forde 
and
Lorenzo 
, 2001;
Weyers 
and
Pater- son 
, 2001). It is frequently observed that the plant’sresponses to drought or P deficiency can be mimicked byexogenouslyapplying a particular plant hormone, though hor-monal involvement in mineral-stress adaptation is discussedcontroversially (
Forde 
and
Lorenzo 
, 2001). But even whenthe root was the stressed organ, hormonal changes havebeen mostly measured in shoot or xylem sap (
Henson 
, 1984;
Itai 
, 1999;
Kozlowski 
and
Pallardy 
, 2002;
Tuberosa 
et al.,1992;
Zhang 
and
Davies 
, 1987). Therefore, the contributionof hormones to drought and phosphorus-deficiency respon-ses of roots was reviewed here.
2 Drought stress
While accumulation of osmolytes like proline or sugars isinduced by severe drought stress, slight changes in plantwater potential may result in a substantial increase in ABAcontent, making this compound a very sensitive drought-stress indicator (
Hartung 
, 1996). The root is a significantsource for ABA in intact plants (
Wilkinson 
and
Davies 
, 2002;
Zhang 
and
Davies 
, 1987). Its biosynthesis in roots and trans-port
via 
the xylem to the shoot may induce drought-stress-adaptation responses in aboveground organs before sub-stantial changes in shoot water potential are detectable.Although it is known that carotenoid precursors of ABA areproduced in plastids and the final steps of ABA biosynthesisare believed to be cytosolic, the precise locations of biosyn-thetic enzymes and translocation pathways of various inter-mediates have not been determined (
Seo 
and
Koshiba 
,2002). Two pathways have been proposed for the synthesisof ABA (
Milborrow 
, 2001). In the “direct pathway”, which hasbeen shown to operate in some fungi, ABA is derived fromfarnesyl diphosphate (
Hirai 
et al., 2000). Because of struc-tural similarities, an “indirect pathway”, in which ABA isproduced from the cleavage of carotenoids, also has beenproposed (
Taylor 
and
Smith 
, 1967). The latter is supported byexperiments with mutant plants investigating genes andenzymatic activities (
Cheng 
et al., 2002;
González-Guzmán 
et al., 2002;
Schwartz 
et al., 2003).Over the last decade, much attention has been paid to ABAconjugates in the xylem sap of various plants (
Sauter 
et al.,2002).
Munns 
and
King 
(1988) and
Munns 
et al. (1993)reported that an unidentified ABA conjugate is more impor-tant as a long-distance stress signal than free ABA. After arri-val in the leaf tissue, this complex form is believed to be phy-siologically more active than ABA.
Bano 
et al. (1993, 1994)were the first to identify ABA glucose ester (ABA-GE) in thexylem sap of rice and sunflower plants. According to
Sauter 
and
Hartung 
(2000) and
Sauter 
et al. (2002), endogenousABA conjugates are formed in the cytosol of root cells, trans-ported symplastically to the xylem parenchyma cells andreleased to the xylem vessels. Because of its extremelyhydrophilic properties, ABA glucose ester (ABA-GE) is trans-located in the xylem of the stem without any loss to the sur-rounding parenchyma. After arrival in the leaf apoplast, trans-porters for ABA-GE in the plasmalemma have to be postu-lated to redistribute the conjugates to the mesophyll cells.Additionally, apoplastic esterases can cleave the conjugateand release free ABA to the target cells and tissues. Theactivity of these esterases is increased when barley plantsare subjected to salt stress.
Wittenmayer 
and
Schilling 
(1998)observed a simultaneous drought-stress-induced increase infree- and conjugated-ABA content of sugar-beet leaves of
2005 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim532 Wittenmayer, Merbach
J. Plant Nutr. Soil Sci.
2005,
168,
531–540
 
intact plants, though sugar-beet tap roots seemingly fail toproduce ABA (
Schulte-Altedorneburg 
et al., 1989).Experiments with the application of free ABA to well-wateredplants support its role in drought-stress adaptation (
Xu 
et al.,2002). The addition of ABA (0.1mmol L
–1
) to the nutrientsolution decreased the average leaf number, area of fullydeveloped leaf and the dry weight per plant of six
Trifolium subterraneum 
L. cultivars by 50%, whereas root : shoot ratiowas increased by 80%. In sugar-beet plants (
Beta vulgaris 
ssp.
vulgaris 
var.
altissima 
Döll), moderate drought stressand application of a nutrient solution containing 0.1mmol L
–1
ABA to well-watered sugar-beet plants resulted in similargrowth responses: reduction of leaf area by 40% andincrease of root : shoot ratio by about 38% (
Wittenmayer 
,2001;
Wittenmayer 
et al., 2004). In this particular crop plant,an ABA-induced increase in root : shoot ratio correlates withthe harvest index (HI) and may contribute to a higher or morestable yield performance under drought stress. In other crops(
i.e.
, where the root does not contribute to the HI), increasedABA content may result in lower yields (
Blum 
et al., 1997).
Saab 
et al. (1990) observed that ABA is necessary for main-taining root growth in drying soil. Later, it has been reportedthat ABA plays a role in promoting root growth independentof its effects on plant water balance (
Sharp 
et al., 2000;
Spol- len 
et al., 2000). It was suggested (
Cheng 
et al., 2002;
Wit- tenmayer 
, 1992) that ABA may possess dual functions: as agrowth inhibitor in the presence of severe drought stress(characterized by induced high endogenous ABA content)and a promoter of root growth in the absence of stress orunder moderate-stress conditions (associated with relativelylow endogenous ABA levels). Alternatively, ABA could func-tion as a growth inhibitor in certain cells, tissues, or organs(
e.g.
, shoot) and as growth promoter in other organs (
e.g.
,roots). The ability of ABA to act as either a promoter or aninhibitor of downstream processes likely depends on thedownstream signaling components, such as transcription fac-tors, that may operate in tissue, development, and/or on theinteractions of ABA with other hormones, such as ethylene(
Eckardt 
, 2002;
Sharp 
, 2002;
Sharp 
and
LeNoble 
, 2002). Forexample, studies in maize (
Spollen 
et al., 2000) suggest thatthe stunted growth of ABA-deficient plants is caused by theoverproduction of ethylene and that one function of ABA maybe to prevent the overproduction of ethylene. Recent resultsindicate a participation in the regulation of aquaporins (
Luu 
and
Maurel 
, 2005).Abscisic acid affects also the accumulation and transport ofosmotically active molecules and ions (
e.g.
, proline, glycinebetaine, Na
+
, K
+
, Ca
2+
, and Cl
) and, thus, the nutritional sta-tus of the plant (
Chen 
and
Murata 
, 2002;
Younis 
et al., 1994).
Younis 
et al. (1994) observed that the addition of ABA at aconcentration of 1mmol L
–1
increased proline and phos-phorus concentration in
Phaseolus vulgaris 
L. Furthermore,abscisic acid stimulates the transport of K
+
, Ca
2+
, and Cl
from root to shoot and accumulation of K
+
, Na
+
, and Cl
inroot cells and inhibits the transport of Na
+
and accumulationof Ca
2+
. It also affects nitrate reductase activity in
Cichorium intybus 
L. (
Goupil 
et al., 1998).Occurrence of ABA is not restricted to plant tissues. The hor-mone occurs also in the rhizosphere and in soil after beingexuded by plant roots or originated by microorganisms (
Hall 
et al., 1986;
Hartung 
et al., 1996;
Müller 
et al., 1989;
Rivier 
etal., 1983;
Slovik 
et al., 1995).
Degenhardt 
et al. (2000) foundthat in alkaline soil substrates, considerable portions of theABA synthesized in the roots leached out into the soil solutionof the rhizosphere.
Passioura 
(2002) speculates that the roottips may continually produce a low level of inhibitor, which infavorable soil conditions, but not in unfavorable, disperses bydiffusing into the neighboring soil. If soil dryness or poor con-tact with the soil (loose soil or roots in biopores) inhibits suchdispersion, this putative inhibitor could build up in concentra-tion within the tip and eventually find its way into the tran-spiration stream and thence to the leaves.
3 Phosphorus deficiency
Plants have evolved sophisticated metabolic and develop-mental strategies to conserve inorganic phosphate (P
i
) and tomaximize its acquisition from the rhizosphere when P
i
is limit-ing (
Ticconi 
et al., 2004). Examples for biochemical adapta-tions include bypass reactions of adenylate- and P
i
-depen-dent steps in respiratory pathways (
Plaxton 
and
Carswell 
,1999), enhanced expression of high-affinity P
i
transporters(
Raghothama 
, 1999) and changes of membrane-lipid compo-sition (
Härtel 
and
Benning 
, 2000;
Yu 
et al., 2002) and ofsugars/photosynthates as crucial components in P-deficiencysignal transduction (
Liu 
et al., 2005;
Müller 
et al., 2005).Remodeling of root-system architecture and increased root-hair formation affecting surface, density, and length of rootsto accelerate soil exploration are typical developmentalresponses to low P
i
, because of low P mobility in soil (
López- Bucio 
et al., 2003;
Ma 
et al., 2001;
Williamson 
et al., 2001).Furthermore, an increased assimilate allocation to rootsresults in a higher root : shoot ratio (
Fredeen 
et al., 1989;
Yan 
et al., 1996).
Lynch 
and
Brown 
(2001) report that root sys-tems with enhanced topsoil foraging acquire phosphorusmore efficiently than others of equivalent size. Comparisonsof contrasting genotypes in controlled environments and inthe field show that plants with better topsoil foraging havesuperior phosphorus acquisition and growth in low-phos-phorus soils. It appears that many architectural responses tophosphorus stress may be mediated by plant hormones (
de Groot 
et al., 2003;
Ma 
et al., 2003). Genetic mapping of thesetraits shows that they are quantitatively inherited, but can betagged with QTLs that can be used in plant-breeding pro-grams (
Lynch 
and
Brown 
, 2001). New crop genotypes incor-porating these traits have substantially improved yield in lowphosphorus soils, and are being deployed in Africa and LatinAmerica (
CIAT 
, 1999;
Ortiz-Monasterio 
et al., 2001). Despitenumerous physiologicalstudies on adaptation responses to Pstarvation, little is known about the underlying hormonal pro-cesses. Table 1 shows the effect of plant growth regulatortreatment applied with nutrient solution on the root surface ofmaize dependingon P availability. In untreated control plants,phosphorus deficiency (–P) increased the root surface by17%. The addition of indole-3-acetic acid (IAA), gibberellicacid (GA
3
), and
trans 
-zeatin (t-Z) to the nutrient solution of–P plants increased the root surface by 31%, 44%, and 49%,respectively. Thus, the tested hormones may increase the
2005 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim
J. Plant Nutr. Soil Sci.
2005,
168,
531540 Plant responses to drought and phosphorusdeficiency 533

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