injury guilds would center on the first injury guild requirement as promulgated byHutchins et al. 42 Injury guild species must produce a similar type of injury.Consequently, similar types of injury can be determined by assessing homogeneitiesof photosynthetic responses. This approach has been used to support injury guildstatus in soybean <strong>and</strong> alfalfa.6.3.3.1 Soybean Leaf-mass Consumer Injury GuildBecause several soybean leaf-mass consumers elicit similar physiological impacts onsoybean, they can be placed into the same injury guild. As discussed above, soybeanleaf-mass consumption does not alter photosynthesis in remaining leaf tissue.Canopy gas exchange responses are altered by leaf-mass consumption injury, butthey most likely are similar among pest species. The other requirements for injuryguild membership (discussed above) also are met for soybean leaf-mass consumers,so several species can be placed into the guild, such as soybean looper, Pseudoplusiaincludens, velvetbean caterpillar, Anticarsia gemmatalis, green cloverworm, beetarmyworm, Spodoptera exigua, <strong>and</strong> bean leaf beetle, Ceratoma trifurcata. 42However, not all species that consume soybean leaves can be members of theleaf-mass consumption injury guild. For example, Mexican bean beetle larvae <strong>and</strong>adults produce a physiological response (reduced photosynthetic rates) different thanthat produced by members of the injury guild. 26 Therefore, the exclusion of theMexican bean beetle from the injury guild is based on physiological response, notphysical appearance of the injury.6.3.3.2 Alfalfa Stubble Defoliator Injury GuildAlfalfa stubble defoliation occurs after the hay crop is cut <strong>and</strong> the stubble is regrowing.Several insects may defoliate alfalfa stubble, including alfalfa weevil larvae <strong>and</strong>adults, Hypera postica, clover leaf weevil larvae <strong>and</strong> adults, Hypera punctata, <strong>and</strong>variegated cutworm larvae, Peridroma saucia. These insects consume dry matter,delay regrowth initiation <strong>and</strong> subsequent plant maturity, <strong>and</strong> reduce growth rates afterdefoliation. 46Peterson et al. 44 concluded that alfalfa responses to clover leaf weevils, alfalfaweevils, <strong>and</strong> variegated cutworms are similar. The three species are leaf-mass consumers.Research on leaf-mass consumption in alfalfa showed no alterations in photosyntheticrates of remaining tissue. 19 Similarities in gas exchange responses inaddition to similarities in consumption patterns <strong>and</strong> timing of injury meet the requirementsfor placing the three species into a common injury guild in alfalfa. Peterson etal. 44 developed injury equivalencies for the three species, which could then be usedfor multiple-species management guidelines.6.4 FUTURE DIRECTIONSFuture research must emphasize how <strong>and</strong> why changes in gas exchange rates occurin response to biotic stress. This type of research will do more to advance our under-
st<strong>and</strong>ing of plant stress from biotic stress than simply characterizing gas exchangeresponses. Using this approach, insect injury <strong>and</strong> other factors, such as plant competitiveinteractions, plant diseases, mineral stress, <strong>and</strong> moisture stress can be integratedinto a more encompassing view of plant stress at all levels of plantorganization.34, 47–49Advances in plant physiology instrumentation <strong>and</strong> biotechnology will helpdetermine both how <strong>and</strong> why changes in plant gas exchange occur after the initiationof biotic stress. In recent years, portable photosynthesis systems (infrared CO 2gasanalyzers) have been developed that allow light response <strong>and</strong> CO 2assimilationcurves to be determined more easily. Determining the genetic basis underlying plantphysiology mechanisms to biotic stress clearly would have far-reaching consequences.Plant transgenic approaches could be employed to better underst<strong>and</strong> <strong>and</strong>manage biotic stress. For example, whole plants <strong>and</strong> canopies of some species, suchas soybean, respond to leaf-mass consumption injury by delaying normal progressivephotosynthetic senescence. If there is a genetic basis for this phenomenon, transgenictechniques could be utilized that would result in greater delays of photosyntheticsenescence after injury, thus tolerating injury better than st<strong>and</strong>ard cultivars.Chapin 50 promulgated a conceptual integration for abiotic stresses. With theappropriate research objectives, a conceptual integration for biotic stresses also ispossible. Finally, integration of abiotic <strong>and</strong> biotic stress will lead to a synthesis for alltypes of plant stress. This will have tremendous value for our underst<strong>and</strong>ing ofnatural <strong>and</strong> agricultural ecosystems.REFERENCES1. Welter, S. C., Arthropod impact <strong>and</strong> plant gas exchange, in Insect-plant Interactions, Vol.1, Bernays, E. A., Ed., CRC Press, Boca Raton, 135, 1989.2. Peterson, R. K. D., <strong>and</strong> Higley, L. G., Arthropod injury <strong>and</strong> plant gas exchange: currentunderst<strong>and</strong>ing <strong>and</strong> approaches for synthesis, Entomol. (Trends Agric. Sci.), 1, 93, 1993.3. Neales, T. F., <strong>and</strong> Incoll, L. D., The control of leaf photosynthesis rate by the level ofassimilate concentration in the leaf: a review of the hypothesis, Bot. Rev., 34, 107, 1968.4. Wareing, P. F., Khalifa, M. M., <strong>and</strong> Treharne, K. J., Rate-limiting processes in photosynthesisat saturating light intensities, Nature, 220, 453, 1968.5. Gifford, R. M., <strong>and</strong> Marshall, C., Photosynthesis <strong>and</strong> assimilate distribution in Loliummultiflorum Lam. following differential tiller defoliation, Aust. J. Biol. Sci., 26, 517,1973.6. Caldwell, M. M., Richards, J. H., Johnson, D. A., <strong>and</strong> Dzurec, R. S., Coping with herbivory:photosynthetic capacity <strong>and</strong> resource allocation in two semiarid Agropyron bunchgrass,Oecologia, 50, 14, 1981.7. Nowak, R. S., <strong>and</strong> Caldwell, M. M., A test of compensatory photosynthesis in the field:implications for herbivore tolerance, Oecologia, 61, 311, 1984.8. Alderfelder, R. G., <strong>and</strong> Eagles, C. F., The effect of partial defoliation on the growth <strong>and</strong>photosynthetic efficiency of bean leaves, Bot. Gaz., 137, 351, 1976.9. Hall, F. R., <strong>and</strong> Ferree, D. C., Effects of insect injury simulation on photosynthesis ofapple leaves, J. Econ. Entomol., 69, 245, 1976.10. Li, J., <strong>and</strong> Proctor, T. A., Simulated pest effects [sic] photosynthesis <strong>and</strong> transpiration of
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Biotic Stressand Yield Loss
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Library of Congress Cataloging-in-P
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PrefaceThe idea for this book came
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EditorsRobert K. D. Peterson, Ph.D.
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ContentsChapter 1Illuminating the B
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1Illuminating the Black Box:The Rel
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increase plant tolerance, through p
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the action of a stressor on a plant
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The magnitude and duration of injur
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Plant part injuredrefers to the pla
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cific competition, while agricultur
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2Yield Loss and PestManagementLeon
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direct relationships between the ac
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In keeping with the theme of this b
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egressions. Actually, the title “
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REFERENCES1. Teng, P. S., Crop Loss
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3Techniques for EvaluatingYield Los
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number of species and stage of cutw
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especially if buried in soil, can d
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elationships for some pests. When m
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injury can be precisely controlled
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day. 81, 99 However, except for an
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the literature most likely are actu
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20. Ba-Angood, S. A., and Stewart,
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60. Stewart, J. G., McRae, K. B., a
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- Page 54 and 55: 4.3.3.1.3 Third generation European
- Page 56 and 57: ing on the developmental stage at t
- Page 58 and 59: 4.2.2.1.2 Temperature stressPlant s
- Page 60 and 61: chronic injury. Acute injury result
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- Page 64 and 65: numbers are present. Stink bugs, Eu
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- Page 82 and 83: 5Phenological Disruptionand Yield L
- Page 84 and 85: ity by animal consumers is the agro
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- Page 88 and 89: FIGURE 5.2 Generalized alfalfa grow
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- Page 106 and 107: 7The Influence of Cultivarand Plant
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- Page 120 and 121: 19. Jarosik, V., Phytoseiulus persi
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- Page 124 and 125: 8Drought Stress, Insects,and Yield
- Page 126 and 127: humidity. Because the relative humi
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- Page 132 and 133: Changes in plant hormones, such as
- Page 134 and 135: plays a key role in promoting plant
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- Page 144 and 145: conditions of stress are themselves
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compensatory response. Also, more v
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Costa Rica, and there are several g
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ivory from white cabbage butterfly
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made, while larger vertebrate herbi
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important consequences to plant fit
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de Entomol., 38, 421, 1994.32. Kare
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chlorophyll content in spider mite
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114. Karban, R., and Strauss, S.Y.,
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10Stephen C. WelterCONTENTSContrast
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Although literature is drawn from a
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and wheat acres receiving some type
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pattern to be true. 109 Because rel
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used in the experiment influenced t
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artificially elevated nitrogen leve
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annual, landrace cultivars, or mode
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settings are coupled with genotype
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10. Kennedy, G. G., and Barbour, J.
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53. Panda, N., and Heinrichs, E. A.
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97. Gross, K. L., and Soule, J. D.,
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143. Davidson, J. L., and Milthorpe
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11Crop Disease andYield LossBrian D
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The conditions listed above are opt
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to associate the effects of disease
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general relationship between LAI an
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Biomassproduction(total dryweight)R
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Y RUE(t)RI(t)[1 X]dt [11.12]wher
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sue. The most accurate prediction o
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tion. Two weeks before harvest, the
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15. Spitters, C. J. T., Van Roermun
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57. Richardson, A. J., Wiegand, C.
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they were cheap, convenient, and ef
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dW / W dtcauses and consequences of
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(a)(b)Maize yield (Mg ha -1 )987654
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Recall that c is a constant, so by
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where the subscripts c and w repres
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0.6Fraction yield loss0.40.2Eq. 16,
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the leaf area index (LAI). Incorpor
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can no longer be tolerated and, the
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cide. Steckel et al. 68 showed that
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A eq ∑ jN eq,ji 1YL n,j [12.31]1
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samples per field. Thomas 85 sugges
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external factors such as annual wea
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38. Boznic, A. C., and Swanton, C.
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weeds, Weed Sci., 44, 856, 1996.79.
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competition and weed management. 3-
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per unit biomass (1/W i)(dW i/dt) o
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of light interception). Algorithms
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where G a,iis the water limited pla
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13.4 COMPETITION FOR SOIL NITROGENA
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As with soil water, Equations 13.10
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partitioning of nitrogen to leaves.
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and stems to optimize photosyntheti
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influence of enhanced UV-B conditio
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Systems Approaches at the Field Lev