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III. Enhancement Strategy for Multiple-Stress Resistance

III. Enhancement Strategy for Multiple-Stress Resistance

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dicates that different mechanisms or degrees of expression in each mechanism are


Specific resistance mechanisms must be associated with adaptive genes. Once

the gene-mechanism relationship is established within a species, this can be used

to identify ecotypes with this resistance mechanism. Gene expression of the mechanism often requires that the abiotidedaphic stresses be present. Also, once a specific stress mechanism is clarified, a search of the scientific literature may reveal

that the associated genes have been identified in other plant species-one of the

objectives of this review article is to assess the current status of gene identification for various abiotic and edaphic stresses. Thus, this component of an overall

strategic framework aids in focusing biotechnology methods and manipulaton on

high-priority genes that are responsible for the specific mechanism@)that causes

a high level of stress resistance.

A third component is to integrate laboratory-generated biotechnology stressenhanced germplasm back into the traditional breeding and genetics programs for

turfgrass improvement. This includes evaluation of germplasm performance under severely stressed and nonstressed field conditions. The following are reasons

for using this approach: (i) Unless the “enhanced germplasm” is placed under the

stress in question, success cannot be determined; (ii) germplasm that has been enhanced for a specific stress mechanism must still perform under multiple stresses;

and (iii) the germplasm can be incorporated into other germplasm by conventional breeding to enhance a broader germplasm pool within a species. Therefore, this

third component integrates biotechnology methods with traditional breeding and

genetic protocols for the purpose of maximizing the efficiency and magnitude of

progress toward greater turfgrass stress performance.




The common abiotic and edaphic stresses that affect turfgrass stress response

plasticity and therefore persistence include

Moisture deficiency/excess problems

Extreme soil/air temperatures

Salinities/poor water quality


High soil strength/traffic/compaction

Low soil oxygen

Reduced light intensity/shade

Low nutrient availability

Many of these constraints interact to create significantgenotype-environment interactions and directly influence turf quality and performance. Turfgrass stress re-



sistance can involve several mechanismsthat may be exhibited at the whole plant to

subcellular level. Each mechanism is genetic based. Most turfgrasses are grown in

multiple-stress (drought-high temperature and soil acidity-drought) environments.

The common plant feature that links all these constraints together is the root system.

The first line of defense in adaptation to abiotic and edaphic stresses is the root

system, which provides essential nutrients and water (i.e., drought resistance) for

critical turfgrass functions. Unfortunately, most turf breeding programs do not address root plasticity (functional root volume and viability under cyclic stresses) either directly or indirectly. Root system improvement should be the first step in

a comprehensive abiotic/edaphic stress-resistancebreeding program because

this strategy addresses primary components of stress response that directly in fluence the turf plant capability to acquire essential nutrients and water and to ultimately persist. Many diverse field situations limit turfgrass rooting, but only six

primary soil chemical and physical constraints account for restricted rooting in

turfgrasses in these field stress situations (Table XIII). These primary stresses can

be incorporated into breeding programs as either single- or multiple-stress screening protocols (Carrow and Duncan, 1996; Duncan and Carrow, 1997; Maranville,

1993). Gene technology can be integrated with this traditional breeding strategy

to enhance genetic-based root plasticity; discern multiple stress tolerance mechanisms; locate, sequence,clone, and map stress-responsivegenes; and utilize marker-assisted selection techniques.

1. Genetic Potential for Rooting

Turf species vary in their genetic potential for rooting depth. Table XIV compares several cool- and warm-season grasses for general root depth potential.

Genotypes within a species can also exhibit inherent differences in rooting depth

under nonlimiting soil conditions (Lehman and Engelke, 1991), but rooting depth

potential is only one component of the overall rhizosphere stress adaptation response mechanism because multiple abiotic and edaphic stresses often limit maximum rooting depth.

While root morphology is governed by genetics (Aeschbacher et al., 1994), developmental plasticity in response to environmental stimuli (light, nutrients, temperature, aeration, water, physical barriers, microorganisms, gravity, competition

from adjacent roots, and chemical barriers) (Schiefelbein and Benfey, 1991) will

ultimately determine the final configuration of the root system (Fitter and Stickland, 1992;Lynch, 1995; Schiefelbein et al., 1997).Developmental alterations occur in the form of changes in the direction of growth after perception of an external signal, transduction of the signal, alteration in gene regulation and protein activity, and modification of cell division-expansion-differentiation (Aeschbacher

et al., 1994). Turf species differ in their capacity for enhanced root growth and

rapid root water uptake at deeper soil layers, maintenance of root viability at the



Table XIII

Six Primary Soil Physical and Chemical Constraintsof Rooting, Associated Field Situations in

Which the StressesAre Expressed, and Relative Importance of Stress on 'hrfgrasses

Relative importance of stress

Root stress

Associated field problems

Fine-textured soil


Layers with few macropores

Sodic soil

Soil drought

Low soil 0,

Fine-textured soils


High water table

Poor surface drainage

Layer impeding percolation

Sodic soil

Acid soil root toxicities (T)/ Acid soil complex (T, D)

deficiencies (D)

AllMnlH toxicities

Nutrient deficiencies (Ca, Mg, K)

Usually low organic matter

Usually high soil strength

Moderately acid soil (D)

Acid sulfate soil (T.D)

Acid mine spoils (T,D)

Sodic soil (T, D)

Salt root toxicities/

Na, CI, B, OH toxicities


K, Mg, Ca deficiencies

High soil strength

Low soil 0,

Saline soil (T, D)

Saline-sodic soil (T, D)

Soil drying


Direct high-temperature root injury

High soil temperature

Indirect high-temperatureb stress

limits root development,

maintenance, viability

High soil strength

Warm season

Cool season













"High organic matter alleviates the Al toxicity factor.

"Indirect high-temperature stress is the major factor limiting cool-season grass adaptation into

warmer temperature climatic zones because it determines carbohydrate status for maintaining root viability. It becomes a site-specific problem when site conditions inhibit canopy cooling. High root temperatures enhance indirect high-temperature stress just as high aerial temperatures will do so.

"The more Xs, the greater the importance.


2 79

lsble XIV

Genetic Potential for Rooting Depth among lbrfgrasses"

Q p e of grass

Root depth


Cool season

Creeping bentgrass

Kentucky bluegrass

Perennial ryegrass

Tall fescue

Warm season

Buffalo grass

Meyer zoysia,

Common centipede

Argentine Bahia grass

Seashore paspalurn (Adalayd)

Emerald zoysia

Tifway Bermuda

Common St. Augustine

Texturf 10 Bermuda

"Reproduced with permission from C m o w (1989).

surface drying layer, and rapid root regeneration after rewatering under drought

conditions (Huang et af., 1997). Roots vary morphologically and physiologically

in response to variable soil nutrient distributions (Robinson, 1996) and to mechanical impedants such as high soil bulk densities or compacted layers (Bengough and Young, 1993; Carrow and Petrovic, 1992; Materechera et af., 1992;

Wiecko et al., 1993). Heritability estimates are quite variable, depending on growing conditions during evaluation and species differences (Browning er af., 1994;

Lehman and Engelke, 1991).

2. Biotechnology

Many of the edaphic and abiotic stress-responsive genes involve the root system. QTLs linked to root morphological characters (Champoux et af., 1995) and

root penetration ability into compacted soils (Ray et af., 1996) have been identified, with potential application in turfgrass transformation studies (Table XV). Increased root density and depth provide an avoidance mechanism in response to

abiotic stress, particularly drought (O'Toole and De Datta, 1986).Compacted soil

layers can impede depth of rooting and negatively enhance the overall stress response in turf. Root penetration ability varies both interspecifically (Assaeed ef al.,

1990; Materechera et af., 1992) and intraspecifically (Kasperbauer and Busscher,

199I ; Masle, 1992) in plants. Screening systems are available to effectively measure root penetration variability among genotypes (Huang et al., 1997; Yu ef aZ.,

1995) and select in the field for root plasticity under stress (Duncan and Carrow,

1997; Erb, 1993; Montpetit and Coulman, 1991).Transgenes governing root gen-



Table XV

QTLs L i e d to Root System Enhancement in Stressed Environments

Root trait


Root thickness, rootshoot ratio, root dry weight per

tiller, deep root dry weight per tiller, maximum root


Root penetration ability into compacted soil layers

Other root-inducing genes (plasmid root-inducing)

rol (root loci: A, B, C, D)

am (auxin synthetic: 1,2)

Champoux ef al. (1995)

Ray et al. ( 1996)

Chriqui er al. (1996)

eration and growth have been expressed in plants (Chriqui et al., 1996). Several

RFLP probes are available to screen for root elongation growth and drought tolerance (Price and Tomos, 1994).Alteration of leaf cytosolic pyruvate kinase can affect source-sink relationships as well as root biomass (Knowles et al., 1998).



While various abiotic and edaphic stresses and their related mechanisms are

genetically controlled, long-term management strategies and variable climatic/

growth conditions will govern turfgrass quality, performance, and persistence.

Most management practices are conducted to alleviate or prevent specific stresses or constraints. Because of the three-way interactions between turf species and

cultivar, specific multiple stresses, and the environment, management strategies

must be adjusted to site-specific situations. Managing the turfgrass root system for

maximum development (depth, volume, and plasticity) and viability/functionality is the key to maintaining high-quality turf in stress environments.

1. Root Management

Root systems in perennial turfgrasses are dynamic or ever-changing (Fig. 3).

Seasonal weather patterns govern growth cycles and affect root topology (branching capacity), root distribution (total biomass, which includes root length and

depth of penetration into the soil), and functionality of roots (root dieback). Most

turf roots survive from 6 months to 2 years, depending on species, management

conditions, and environmental constraints (Carrow, 1989). Duration of exposure

and severity of a stress or multiple stresses have profound influences on turfgrass

persistence mainly because carbohydrates produced in green shoot tissues by pho-



28 1


Figure 3 Seasonal root growth rates of turfgrasses.

tosynthesis are usually utilized first for shoot growth and maintenance and secondarily for root growth and maintenance. Severe environmental stress will create

unbalanced carbohydrate demands in turf plants that can enhance root mortality or

decrease root functionality and ultimately diminish turf quality.

Proper root management to minimize stress is essential to turf quality longevity (Carrow, 1989, 1995a):

1. Select species and cultivars within species with the best root plasticity capability.

2. Promote maximum net carbohydrate production by

a. Optimizing leaf area, which ensures maximum photosynthesis, by increasing mowing height, decreasing wear damage, and controlling biotic


b. Optimizing leaf chlorophyll content by avoiding (i) Fe, Mn, Mg, S, and

M deficiencies, (ii) low soil oxygen or waterlogged conditions, and

(iii) prolonged water-deficit conditions.

c. Promoting good light capture conditions by (i) pruning trees and removing excess grass clippings and (ii) selecting appropriate cultivars.

3. Avoid depletion of carbohydrate reserves in the crown region by minimizing excessive and frequent N applications (especially fast-release N sources),

overwatering, and close mowing. Modify high soil temperatures that contribute to

the depletion of carbohydrates with imgation, drainage, cultivation, or by increasing mowing height.

2 82


4. Correct soil physical problems as follows: Correct high soil strength (i.e.,

high bulk density and heavy clay soils) and low soil oxygen with cultivation (aeration) and additions of peat or gypsum; excessively dry soils with irrigation and

additions of organic matter to increase water-holding capacity; low soil oxygen

with cultivation and surface/subsurface drainage; soil layering with cultivation;

and cold soils in the spring with cultivation and proper drainage.

5. Correct poor soil chemical conditions as follows: Correct acid/high Al soils

with lime; very alkaline soils with S, H,SO,, or acidic N carriers; infertile soils

with fertilizers or microbial amendments; and salt-affected soils with cultivation,

gypsum, or sulfur amendments, drainage, and use of alternative water sources.

Avoid toxins by limiting excessive use of herbicides or other chemicals, limiting

heavy metal-containing soil amendments, and judicious application of macro- and


6. Correct soil biotic problems as follows: Correct root-feeding insects, diseases, and nematodes with preventive, cultural, or chemical control treatments;

thatch by mechanical removal, cultivation, and promotion of microbial degradation.


Perennial grasses will always be subjected to fluctuating multiple stresses. Traditional breeding programs can address specific environmental constraints and, as

mechanisms governing stress response become better understood, these programs

can focus on specific components of these mechanisms. Gene technology provides

an enhancement strategy for these traditional breeding approaches. An increasing

number of genes are being identified, sequenced, and cloned. Transformation and

regeneration technology is available for implementation into turfgrass stressresistance programs. With the release of new “biotech” turf cultivars in the twenty-first century, management strategies will have to be adjusted to maximize performance and persistence.

Enhanced abiotidedaphic stress tolerance in turf will provide

1. Improvements in performance under environmental extremes

2. Functional root systems that perform equally well in stressed and nonstressed environments

3. Improved water use efficiency

4. Improved nutrient uptake/utilization efficiency

5. Better adapted cultivars for niche environments

6. More high-quality and environmentally compatible turfgrasses under abiotic/edaphic stressed conditions




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