<str<strong>on</strong>g>Proceedings</str<strong>on</strong>g> <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> <str<strong>on</strong>g>Third</str<strong>on</strong>g> <str<strong>on</strong>g>Internati<strong>on</strong>al</str<strong>on</strong>g> <str<strong>on</strong>g>C<strong>on</strong>ference</str<strong>on</strong>g> <strong>on</strong> <strong>Invasive</strong> SpartinaChapter 1: Spartina BiologyFig. 4. Alcohol dehydrogenase (ADH) activities (μmol g -1 min -1 ) <str<strong>on</strong>g>of</str<strong>on</strong>g>Spartina and Distichlis grown under drained and flooded soil treatments.Shown is <str<strong>on</strong>g>the</str<strong>on</strong>g> mean <str<strong>on</strong>g>of</str<strong>on</strong>g> 7-13 plants ± SE. Species are labeled as in Fig. 2.activities in S. anglica were significantly lower than all o<str<strong>on</strong>g>the</str<strong>on</strong>g>rspecies (ANOVA, p≤0.046). Flooded soil c<strong>on</strong>diti<strong>on</strong>sresulted in significantly higher ADH activities in all species(ANOVA, p≤0.018) expect <str<strong>on</strong>g>the</str<strong>on</strong>g> low marsh species S. anglica(ANOVA, p=0.564).Most plants can survive short term absence <str<strong>on</strong>g>of</str<strong>on</strong>g> oxygen(≤60 min) without cell death. In <str<strong>on</strong>g>the</str<strong>on</strong>g>se cases, normal ATPstores are quickly depleted in active cells, and mitoch<strong>on</strong>drialswelling is usually observed within minutes (Drew 1997).Large reserves <str<strong>on</strong>g>of</str<strong>on</strong>g> stored carb<strong>on</strong> can help <str<strong>on</strong>g>the</str<strong>on</strong>g> cells respireanaerobically, but l<strong>on</strong>ger absences <str<strong>on</strong>g>of</str<strong>on</strong>g> oxygen can result incell death. Irreversible damage to mitoch<strong>on</strong>dria and cellviability generally occurs following 15 hours <str<strong>on</strong>g>of</str<strong>on</strong>g> anaerobiosis(Perata and Alpi 1993). However, a supply <str<strong>on</strong>g>of</str<strong>on</strong>g> oxygen frominternal aerati<strong>on</strong> can allow marsh plants like Spartina torespire aerobically despite growing in waterloggedsubstrates. The superior oxygen transport abilities <str<strong>on</strong>g>of</str<strong>on</strong>g> S.anglica may have helped to account for its low root ADHactivities observed in this study.The plants studied showed varying degrees <str<strong>on</strong>g>of</str<strong>on</strong>g> sulfideoxidati<strong>on</strong> capacity. Total sulfide oxidati<strong>on</strong> was partiti<strong>on</strong>edinto enzymatic and n<strong>on</strong>enzymatic processes. The meanenzymatic sulfide oxidase (SOx) activity ranged from 14.4to 97.2 nmol g -1 min -1 across species and waterloggingtreatments (Fig. 5a). Enzymatic SOx activities were highestin <str<strong>on</strong>g>the</str<strong>on</strong>g> high marsh species S. patens and D. spicata and weresignificantly lower in S. densiflora and <str<strong>on</strong>g>the</str<strong>on</strong>g> low marshspecies S. alterniflora and S. anglica (ANOVA, p≤0.050).This difference may suggest that high marsh species aremore sensitive to sediment sulfides and thus require greaterenzymatic protecti<strong>on</strong>. Enzymatic SOx activities did notchange in resp<strong>on</strong>se to flooding across species (ANOVA,p≥0.960).N<strong>on</strong>biotic factors such as metal i<strong>on</strong>s can c<strong>on</strong>tribute tosulfide oxidati<strong>on</strong> (Lee et al. 1999). Such n<strong>on</strong>enzymaticprocesses were also found to be important in sulfideoxidati<strong>on</strong> in <str<strong>on</strong>g>the</str<strong>on</strong>g> present study. Mean n<strong>on</strong>enzymatic sulfideFig. 5. Sulfide oxidase (SOx) activities (nmol g -1 min -1 ) <str<strong>on</strong>g>of</str<strong>on</strong>g> Spartina andDistichlis grown under drained and flooded soil treatments. Shown are (a.)enzymatic and (b.) n<strong>on</strong>enzymatic rates <str<strong>on</strong>g>of</str<strong>on</strong>g> sulfide oxidati<strong>on</strong>. The mean <str<strong>on</strong>g>of</str<strong>on</strong>g>4-10 plants are shown ± SE. Species are labeled as in Fig. 2.oxidati<strong>on</strong> rates ranged from 12.6 to 51.1 nmol g -1 min -1across species and waterlogging treatments (Fig. 5b).N<strong>on</strong>enzymatic rates <str<strong>on</strong>g>of</str<strong>on</strong>g> sulfide oxidati<strong>on</strong> were not differentbetween species or flooding treatment (ANOVA, p≥0.141).CONCLUSIONSThe upper regi<strong>on</strong>s <str<strong>on</strong>g>of</str<strong>on</strong>g> salt marshes are characterized byoxidized soils, since tidal flooding is rare. However, episodicflooding at <str<strong>on</strong>g>the</str<strong>on</strong>g> highest tides can result in occasi<strong>on</strong>al anoxicand sulfidic c<strong>on</strong>diti<strong>on</strong>s. Therefore, plants from <str<strong>on</strong>g>the</str<strong>on</strong>g> highmarsh are not forced to withstand chr<strong>on</strong>ic anoxia. The highmarsh species S. patens, S. densiflora, and Distichlis spicatawere found to have high aerobic respirati<strong>on</strong> rates and highaerobic enzyme activity. This aerobic oxygen demand maybe too high to allow survival in anoxic low marshc<strong>on</strong>diti<strong>on</strong>s, where plants had lower aerobic demand.Anaerobic pathways (root ADH activities) increased afterflooding in all three high marsh species suggesting a highsensitivity to soil waterlogging. Internal oxygen transportrates were low in <str<strong>on</strong>g>the</str<strong>on</strong>g>se plants since <str<strong>on</strong>g>the</str<strong>on</strong>g>y are adapted to lifein sediments where soil oxygen is not normally limiting toroot respirati<strong>on</strong>.Additi<strong>on</strong>ally, higher SOx activities were found in highmarsh species compared to low marsh species. These trendssuggested that high marsh species were more sensitive tosulfide and required greater protecti<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> aerobic respirati<strong>on</strong>.This idea is c<strong>on</strong>sistent with <str<strong>on</strong>g>the</str<strong>on</strong>g> finding that <str<strong>on</strong>g>the</str<strong>on</strong>g>se speciesexhibited higher activities <str<strong>on</strong>g>of</str<strong>on</strong>g> CytOx, <str<strong>on</strong>g>the</str<strong>on</strong>g> site <str<strong>on</strong>g>of</str<strong>on</strong>g> sulfideinhibiti<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> aerobic respirati<strong>on</strong> (Bagarinao 1992). Highrates <str<strong>on</strong>g>of</str<strong>on</strong>g> aerobic respirati<strong>on</strong> and apparent sulfide sensitivitymay substantially account for <str<strong>on</strong>g>the</str<strong>on</strong>g> exclusi<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g>se speciesfrom low marsh z<strong>on</strong>es.The low z<strong>on</strong>es <str<strong>on</strong>g>of</str<strong>on</strong>g> salt marshes are characterized byfrequent tidal flooding. This leads to highly reducedsediments, <str<strong>on</strong>g>of</str<strong>on</strong>g>ten c<strong>on</strong>taining high levels <str<strong>on</strong>g>of</str<strong>on</strong>g> sulfides. Plantsinhabiting low marsh regi<strong>on</strong>s must be able to tolerate highlyreducing and sulfidic sediment c<strong>on</strong>diti<strong>on</strong>s. Spartinaalterniflora is <str<strong>on</strong>g>the</str<strong>on</strong>g> dominant low marsh species in manyNorth American East- and Gulf Coast estuaries (Bertness-51-
Chapter 1: Spartina Biology<str<strong>on</strong>g>Proceedings</str<strong>on</strong>g> <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> <str<strong>on</strong>g>Third</str<strong>on</strong>g> <str<strong>on</strong>g>Internati<strong>on</strong>al</str<strong>on</strong>g> <str<strong>on</strong>g>C<strong>on</strong>ference</str<strong>on</strong>g> <strong>on</strong> <strong>Invasive</strong> Spartina1991). Spartina anglica can grow lower in <str<strong>on</strong>g>the</str<strong>on</strong>g> intertidalrange than S. alterniflora (Frenkel 1987, Sayce andMumford 1990), and <str<strong>on</strong>g>the</str<strong>on</strong>g>refore any o<str<strong>on</strong>g>the</str<strong>on</strong>g>r species in thisstudy. Low marsh species exhibited low aerobic respirati<strong>on</strong>rates and CytOx activity, which when coupled to high rates<str<strong>on</strong>g>of</str<strong>on</strong>g> internal oxygen transport may pose a significantadvantage for survival in anoxic sediments. The ability tosupply oxygen to submerged tissue is crucial to survival in<str<strong>on</strong>g>the</str<strong>on</strong>g> low marsh, as no plant tissue can endure anoxiaindefinitely (Crawford 1982). Low marsh species must alsopossess an ability to respire anaerobically since demand foroxygen from highly reduced sediments may overwhelmtransport processes. ADH activities measured in Spartinaroots indicated a well-developed capacity for fermentati<strong>on</strong>.However, increases in root ADH were not observed in <str<strong>on</strong>g>the</str<strong>on</strong>g>low marsh species S. anglica. High rates <str<strong>on</strong>g>of</str<strong>on</strong>g> oxygen transportin S. anglica may be adequate to supply oxygen to roots andexternal sinks, as suggested by its low root ADH activities.Low marsh species may be more resistant to sulfides whencompared to high marsh species. Lower aerobic respirati<strong>on</strong>rates and lower CytOx activities may relax needs for intensesulfide oxidati<strong>on</strong> requirements. Alternatively, higher rates <str<strong>on</strong>g>of</str<strong>on</strong>g>oxygen transport in some low marsh species may help tooxidize rhizosphere sulfides, resulting in reduced SOxactivity. However, <str<strong>on</strong>g>the</str<strong>on</strong>g> acute toxicity <str<strong>on</strong>g>of</str<strong>on</strong>g> dissolved sulfidesaround roots still necessitates moderate SOx activities in lowmarsh species.The results <str<strong>on</strong>g>of</str<strong>on</strong>g> this study suggest metabolic characteristicsrelated to respirati<strong>on</strong> and sulfide tolerance may affect z<strong>on</strong>ati<strong>on</strong><str<strong>on</strong>g>of</str<strong>on</strong>g> grasses in estuaries. While internal oxygen transport isimportant for survival in estuarine sediments, this study mayindicate that z<strong>on</strong>ati<strong>on</strong> within estuaries is dependent <strong>on</strong> morethan just oxygen transport. Aerobic respirati<strong>on</strong> rates andsensitivity to sediment sulfides may play a large role ininfluencing estuarine z<strong>on</strong>ati<strong>on</strong> as well.ACKNOWLEDGMENTSThe authors thank Chuck Cody for greenhouse assistance;Kim Patten, Sally Hacker, Eric Hellquist, and M. Enrique Figueroafor providing plants; and <str<strong>on</strong>g>the</str<strong>on</strong>g> Washingt<strong>on</strong> State Department <str<strong>on</strong>g>of</str<strong>on</strong>g>Agriculture for a Spartina transport permit. This project waspartially funded from <str<strong>on</strong>g>the</str<strong>on</strong>g> Betty W. Higinbotham Trust, NSFIBN0076604, EPA grant R-82940601, and NSF DBI-0116203.REFERENCESAdam, P. 2002. Saltmarshes in a time <str<strong>on</strong>g>of</str<strong>on</strong>g> change. Envir<strong>on</strong>mentalC<strong>on</strong>servati<strong>on</strong> 29:39-61.Arenovski, A.L., and B.L. Howes. 1992. Lacunal allocati<strong>on</strong> and gastransport capacity in <str<strong>on</strong>g>the</str<strong>on</strong>g> salt marsh grass Spartina alterniflora.Oecologia 90:316-322.Armstr<strong>on</strong>g, W. 1964. Oxygen diffusi<strong>on</strong> from <str<strong>on</strong>g>the</str<strong>on</strong>g> roots <str<strong>on</strong>g>of</str<strong>on</strong>g> someBritish bog plants. Nature 204:801-802.Bagarinao, T. 1992. Sulfide as an envir<strong>on</strong>mental factor and toxicant:Tolerance and adaptati<strong>on</strong>s in aquatic organisms. AquaticToxicology 24:21-62.Bertness, M.D. 1991. Z<strong>on</strong>ati<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> Spartina patens and Spartinaalterniflora in a New England salt marsh. Ecology 72:138-148.Bray, E.A., J. Bailey-Serres, and E. Weretilnyk. 2000. Resp<strong>on</strong>ses toabiotic stresses. In: Buchanan, B. B., W. Gruissem, and R. L.J<strong>on</strong>es, eds. Biochemistry and Molecular Biology <str<strong>on</strong>g>of</str<strong>on</strong>g> Plants.American Society <str<strong>on</strong>g>of</str<strong>on</strong>g> Plant Physiologists: Rockville, Maryland.pp. 1158-1203.Brix, H., and B.K. Sorrell. 1996. Oxygen stress in wetland plants:comparis<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> de-oxygenated and reducing root envir<strong>on</strong>ments.Functi<strong>on</strong>al Ecology 10:521-526.Burdick, D.M., and I.A. Mendelssohn. 1987. Waterlogging resp<strong>on</strong>sesin dune, swale, and marsh populati<strong>on</strong>s <str<strong>on</strong>g>of</str<strong>on</strong>g> Spartina patensunder field c<strong>on</strong>diti<strong>on</strong>s. Oecologia 74:321-329.Byrd, G.T., R.F. Sage, and R.H. Brown. 1992. A comparis<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g>dark respirati<strong>on</strong> between C 3 and C 4 plants. Plant Physiology100:191-198.Cline, J.D. 1969. Spectrophotometric determinati<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> hydrogensulfide in natural waters. Limnology and Oceanography 14:454-458.Crawford, R.M.M. 1967. Alcohol dehydrogenase activity in relati<strong>on</strong>to flooding tolerance in roots. Journal <str<strong>on</strong>g>of</str<strong>on</strong>g> Experimental Botany18:458-464.Crawford, R.M.M. 1982. Physiological resp<strong>on</strong>ses to flooding. In:Lange, O.S. P.S. Nobel, C.B. Osm<strong>on</strong>d, and H. Ziegler, eds. Encyclopedia<str<strong>on</strong>g>of</str<strong>on</strong>g> Plant Physiology, Physiolological Plant EcologyII. Water Relati<strong>on</strong>s and Carb<strong>on</strong> Assimilati<strong>on</strong>. Springer-Verlag:Berlin. pp. 453-477Drew, M.C. 1997. Oxygen deficiency and root metabolism: Injuryand acclimati<strong>on</strong> under hypoxia and anoxia. Annual Review <str<strong>on</strong>g>of</str<strong>on</strong>g>Plant Physiology and Plant Molecular Biology 48:223-250.Emmett, R., R. Llanso, J. Newt<strong>on</strong>, R. Thom, M. Hornberger, C.Morgan, C. Levings, A. Copping, and P. Fishman. 2000. Geographicsignatures <str<strong>on</strong>g>of</str<strong>on</strong>g> North American West Coast estuaries. Estuaries23:765-792.Epstein, E. 1972. Mineral Nutriti<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> Plants: Principles and Perspectives.John Wiley and S<strong>on</strong>s, Inc.: New York.Frenkel, R.E. 1987. Introducti<strong>on</strong> and spread <str<strong>on</strong>g>of</str<strong>on</strong>g> cordgrass (Spartina)into <str<strong>on</strong>g>the</str<strong>on</strong>g> Pacific Northwest. Northwest Envir<strong>on</strong>mental Journal3:152-154.Gleas<strong>on</strong>, M.L., and J.C. Zieman. 1981. Influence <str<strong>on</strong>g>of</str<strong>on</strong>g> tidal inundati<strong>on</strong>in internal oxygen supply <str<strong>on</strong>g>of</str<strong>on</strong>g> Spartina alterniflora andSpartina patens. Estuarine, Coastal and Shelf Science 13:47-57.Hacker, S.D., D. Heimer, C.E. Hellquist, T G. Reeder, B. Reeves,T.J. Riordan, and M.N. Dethier. 2001. A marine plant (Spartinaanglica) invades widely varying habitats: Potential mechanisms<str<strong>on</strong>g>of</str<strong>on</strong>g> invasi<strong>on</strong> and c<strong>on</strong>trol. Biological Invasi<strong>on</strong>s 3:211-217.Hedge, P., L.K. Kriwoken, and K. Patten. 2003. A review <str<strong>on</strong>g>of</str<strong>on</strong>g>Spartina management in Washingt<strong>on</strong> State, US. Journal <str<strong>on</strong>g>of</str<strong>on</strong>g>Aquatic Plant Management 41:82-90.Howes, B.L., and J.M. Teal. 1994. Oxygen loss from Spartina alternifloraand its relati<strong>on</strong>ship to salt marsh oxygen balance.Oecologia 97:431-438.Hwang, Y.-H., and J.T. Morris. 1991. Evidence for hygrometricpressurizati<strong>on</strong> in <str<strong>on</strong>g>the</str<strong>on</strong>g> internal gas space <str<strong>on</strong>g>of</str<strong>on</strong>g> Spartina alterniflora.Plant Physiology 96:166-171.Jacks<strong>on</strong>, M.B., and W. Armstr<strong>on</strong>g. 1999. Formati<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> aerenchymaand <str<strong>on</strong>g>the</str<strong>on</strong>g> process <str<strong>on</strong>g>of</str<strong>on</strong>g> plant ventilati<strong>on</strong> in relati<strong>on</strong> to soil floodingand submergence. Plant Biology 1:274-287.John, C.D., and H. Greenway. 1976. Alcoholic fermentati<strong>on</strong> andactivity <str<strong>on</strong>g>of</str<strong>on</strong>g> some enzymes in rice roots under anaerobiosis. AustralianJournal <str<strong>on</strong>g>of</str<strong>on</strong>g> Plant Physiology 3:325-336.-52-
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