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UNIVERSIDAD CENTRAL DE VENEZUELA COMISIN DE ESTUDIOS PARA GRADUADO POSTGRADO EN AGRONOMA MARACAY

PROGRAMA ASIGNATURA: Fisiologa de herbicidas CDIGO: Profesores asignatura: Dra. Ada Ortiz (Coordinadora) PhD. Albert Fischer (Universidad de California-Davis, EE.UU) Dr. Alvaro Anzalone (Universidad Centroccidental Lisandro Alvarado, Barquisimeto, Lara) CRDITOS: Tres HORAS/SEMANA: 48 horas (Lunes a sbado). AO: 2013 COMPONENTE: Electiva en el Postgrado en agronoma CONOCIMIENTOS DE ENTRADA: 1. 2. 3. 4. 5. Fisiologa Vegetal Ecologa agrcola Bioqumica Manejo integrado de cultivo Ingls instrumental Aprobado por: Postgrado en Agronoma en fecha: Comisin de Estudios de Postgrado de Agronoma en fecha: Consejo de Facultad de Agronoma en fecha:

Naturaleza: Terica Ubicacin: II Perodo Distribucin horaria: 8 horas por da, una semana Horario: diurno Duracin: 48 horas, Asignatura diseada para dictarlo en una semana

JUSTIFICACIN La presencia de malezas en los cultivos causa daos directos e indirectos, los primeros son los producidos por efecto de la interferencia (competencia, parasitismo o alelopata) y los otros porque encarecen la produccin de cultivo y deterioran su calidad, reducen el valor de las fincas y son hospedantes alternos de diversas plagas. El estudio de las malezas y su manejo es parte del eje transversal del currculo de las carreras relacionadas a la agronoma.

PROPSITO En esta asignatura se consideran en forma detallada los principios tericos que rigen el comportamiento de los herbicidas en el suelo, en masas de agua y en la planta, tales como: (a) Comportamiento de los herbicidas en el suelo, procesos de adsorcin, volatilizacin, degradacin y persistencia, (b) Tcnicas de deteccin de residuos de herbicidas en suelo y agua, (c) Absorcin y transporte de herbicida en la planta, (d) Coadyuvantes, (e) Metabolismo de herbicida y selectividad, (f) Antdotos (protectantes), (g) Modo de accin de los herbicidas: Inhibidores de la enzima ALS, inhibidores de la biosntesis de lpidos (inhibidores de las enzimas ACCasa y elongasas); herbicidas que resultan en formas txicas de oxgeno (bipiridilos, inhibidores de las enzimas Protox y HPPD, e inhibidores de la sntesis de carotenoides), inhibidores del fotosistema II, inhibidores de la divisin celular, herbicidas auxnicos, glifosato y glufosinato y (h) Resistencia de malezas a herbicidas y su manejo ORGANIZACIN: 1. Coordinador de Asignatura (Institucin): Ada Ortiz (FAGRO-UCV) 2. Profesores (Institucin): Aida Ortiz (FAGRO-UCV), Albert Fischer (UC-Davis) y lvaro Anzalone (UCLA). 3. Sesiones tericas a dictarse en las aulas disponibles del Postgrado de FAGRO-UCV o la UCLA Recursos disponibles Planta fsica: 1 saln con capacidad para 30 personas Docentes: 3 docentes Recursos audiovisuales: video beam, pizarrn, Sofware libre Recursos computacionales: laboratorio con computadoras ESTRATEGIAS INSTRUCCIONALES Exposicin del docente con medios audiovisuales Lecturas complementarias Dinmicas de Grupo

PROGRAMA SINPTICO EDICIN REVISIN CUATRIMESTRE CDIGO Fisiologa de herbicidas 3 PRELACIONES Presencial HORAS SEMANA

1 II

ASIGNATURA UNIDAD CRDITO MODALIDAD: 48 h OBJETIVO GENERAL: Comprender el funcionamiento de los herbicidas dentro de la planta as como su interaccin con el medio ambiente a fin de afianzar sus conocimientos sobre el manejo integrado de malezas y evitar la evolucin de la resistencia de malezas a herbicidas. OBJETIVOS ESPECFICOS 1. Reconocer el comportamiento de los herbicidas en el suelo y en el agua. 2. Definir algunas tcnicas de deteccin de herbicidas en suelo y agua 3. Relacionar los procesos de absorcin y transporte de los herbicidas hacia los sitios de accin en la planta 4. Seleccionar el coadyuvante ms apropiado para mejorar la absorcin de los herbicidas en la planta 5. Revisar los procesos metablicos de herbicidas en las plantas conducentes a selectividad de cultivos o resistencia de malezas 6. Escoger el antdoto adecuado para mejorar la selectividad de los cultivos a los herbicidas 7. Comprender los modos de accin de los herbicidas para racionalizar su uso 8. Reconocer la resistencia de malezas a herbicidas y sus mecanismos bioqumicos y fisiolgicos 9. Programar estrategias de manejo de la resistencia de malezas a herbicidas SINPTICO DE CONTENIDOS ETAPA 1 1. Herbicidas en suelo y agua: Procesos de adsorcin, volatilizacin, degradacin y persistencia de los herbicidas en el suelo y en el agua 2. Tcnicas de deteccin de residuos de herbicidas en suelo y agua ETAPA 2 1. Absorcin y transporte de herbicidas en la planta 2. Coadyuvantes para mejorar la absorcin de los herbicidas en la planta 3. Metabolismo de herbicidas y selectividad 4. Uso de antdotos para incrementar las selectividad de los cultivos ETAPA 3. Modo de accin de los herbicidas 1. Inhibidores de la enzima ALS 2. Inhibidores de la biosntesis de lpidos (inhibidores de las enzimas ACCase y elongasas) 3. Herbicidas que resultan en formas txicas de oxgeno: a. Bipiridilos b. Inhibidores de las enzimas Protox y HPPD

4. 5. 6. 7. 8.

c. Inhibidores de las sntesis de carotenoides), Inhibidores del fotosistema II Inhibidores de la divisin celular y de la pared celular Herbicidas auxnicos Glifosato Glufosinato

ETAPA 3. Resistencia de malezas a herbicidas y su manejo 1. Definicin de resistencia 2. Mecanismos de resistencias 3. Mitigacin EVALUACIN Se enviar por correo electrnico, a cada estudiante inscrito en esta asignatura, un examen correspondiente a la evaluacin del 100% de la asignatura dictada, el cual tendr una calificacin del 0 a 20 puntos.

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ABSORCIN Y TRANSPORTE DE HERBICIDA EN LA PLANTA 1. Briggs, G.G., R.H. Bromilow, A.A. Evans 1982. Relationships between lipohilicity and root uptake and translocation of non-ionised chemicals by barley. Pesticide Science 13:495-504. 2. Briskin, D. P.; 1994. Membrane and transport system in plants: An overwiew. Weed Science, 42:255-262. Bromilow, R.H., K. Chamberlain, A.A. Evans. 1990. Physicochemical aspects of phloem translocation of herbicides. Weed Science 38:305314. 3. Bromilow, R.H., K. Chamberlain. 2000. The herbicide glyphosate and related molecules: physicochemical and structural factors determining their mobility in phloem. Pest Managernent Science. 56:368-373. 4. Bromilow R., K. Chamberlain and A. Evans. 1990. Physicochemical aspects of phloem translocation of herbicides. Weed Sci. 38:305-314 5. Crafts, A. S. 1961. The Chemistry and Mode of Action of Herbicides. Interscience Publishers, New York, London. Chapter 5. 6. Crafts, A. S., E. C. Crisp. 1971. Phloem transport in plants. Chap. W. H. Freeman: San Francisco, CA. 7. Crisp, C.E. 1972. The molecular design of systemic insecticides and organic functional groups in translocation. In Pesticide Chemistry: Proceedings of the Second International I[JPAC Congress. Vol 1: Insecticides. Tel Aviv, Israel. p 211-264. 8. Crisp, C.E., M. Look. 1979. Phloem loading and transport of week acids. In Advances in Pest Science. Vol 3. Ed by H. Geissbuhler. Pergamon press, New York. pp 430-437. 9. Devine,M.D., H.D. Bestman, W.H. Vanden Born. 1987. Uptake and Accumulation of the herbicides chlorsulfuron and clopyralid in excised pea root tissue. Plant Physiology. 85:82. 10. Devine, M.D., S.O. Duke, and C. Fedtke. Physiology of Herbicide Action. 1993. Prentice Hall, NJ. Pp. 67-94 11. Dewey, S.A. and A.P. Appleby. 1983. A comparison between glyphosate and assimilate translocation patterns in tall morningglory (Ipomoea purpurea). Weed Sci. 31:308-314. 12. Donaldson, T. W., D. E. Bayer, O. A. Leonard. Absorptiion of 2,4Dichlorophenoxyacetic acid an 3-(pChlorophenyl)-1,1-dimetilhylurea (Monuron) by barley roots. Plant Physiology, 52:638-645. 13. Edgington, L. V.; C. A. Peterson. 1977. Systemic fungicides: theory, uptake and translocation. In M.R. Siegel and H.D. Sisler (eds.) Antifungal Compounds, vol. 2, Interactions in Ecological Systems, Marcel Dekker, New York. Pp. 51-89. 14. Geiger D.R., H.D. Bestman. 1990. Self-limitation of herbicide mobility by phytotoxic action. Weed Sci. 38:324329 15. Hsu, F.C., D.A. Kleier. 1996. Phloem mobility of xenobiotics VIII. A short review. Journal of Experimental Botany. 47:1265-1271. 16. Kasai, F, D.E. Bayer. 1991. Quantitative evaluation of the weak acid hypothesis as the mechanism for 2,4-D absorption by corn root protoplasts. Journal of Pesticide Science 16:163. 17. Kleier, D.A. 1988. Phloem mobility of xenobiotics. I. Mathematical model unifying the weak acid and intermediate permeability theories. Plant Physiology. 86:803-810.

18. Kleier, D.A., F.C. Hsu. 1996. Phloem mobility of xenobiotics. VII. The design of phloem systemic pesticides. Weed Science. 44:749-756. 19. Klingman, G.C., F.M. Ashton, L.J Noordhoff. 1975. Weed Science: Principles and Practices 2d ed. John Wiley and Sons, New York, pp 62-67. 20. Palmgreen, M. G. 2001. Plant plasm membrane H+ATPases: powerhouses for nutrient uptake. Annual Review of Plant Physiology and Plant Molecular Biology. 52:817-845. 21. Raven, J.A. 1975. Transport of indoleacetic acid in plants cells in relation to pH [hydrogen ion concentration] and electrical potential gradients, and its significance for polar IAA transport. New Phytol. 74:163. Satchivi, N.M., E.W. Stoller, L.M. Wax, et al. 2001. A nonlinear dynamic simulation model for xenobiotic transport and whole plant allocation following foliar application. III. Influence of chemical properties, plant characteristics, and environmental parameters on xenobiotic absorption and translocation. Pesticide Biochemistry and Physiology. 71:77-87. 22. Smith, F.A., J.A. Raven. 1979. Intracellular pH hydrogen -ion concentration and its regulation Cytoplasm and vacuole of plant cells. Annual. Review of Plant Physiology. 30:289. 23. Sterling, T. M. 1994. Mechanism of herbicide absorption across plant membranes and accumulation in plan cells. Weed Science. 42:263-276. 24. Taiz, L., E. Zeiger. 2002. Plant Physiology. 3 ed. Sinauer Associates, Inc., Publishers, Massachusetts. Chapter 6. 25. Tice, C.M. 2001. Selecting the right compounds for screening: does Lipinski's rule of 5 for pharmaceuticals apply to agrochemicals? Pest Managernent Science. 57:3-16. 26. Theodoulou, F.L. (2000) Plant ABC transporters. Biochim. Biophys. Acta 1465:. 79103. 27. Tyree, M.T., C.A. Peterson, L.V. Edgington. 1979. A simple theory regarding the ambimobility of xenobiotics with special reference to the nematicide, oxamyl. Plant Physiology. 63:367. 28. Weeding, R.T., L.C. Erickson. 1957. The role of pH in the permeability of Chlorella to 2,4-D. Plant Physiology 32:503-512. 29. Xoconostle-Cazares, X.B., Y. Xiang, R.R. Medrano,et al. 1999. Plant paralog to viral movement protein that potentiates transport of mRNA into the phloem. Science. 283:9498. COADYUVANTES 1. Balke and Price. 1988. Pesticide Biochemistry and Physiology 30:228-237. 2. Briskin, D. P. 1994. Membranes and tansport systems in plants: An overview. Weed Sci. 42: 255-262 Donaldson et al. 1973. Plant physiol 52:638-645 3. Devine et al. 1987. Plant Physiol. 85:82-86. 4. Devine, M.D., S.O. Duke, and C. Fedtke. Physiology of Herbicide Action. 1993. Prentice Hall, NJ. Pp. 29-65 5. Hart J. J.; Di Tomaso J. M. ; Linscott D. L. ; Kochian L. V. 1992. Characterization of the transport and cellular compartmentation of paraquat in roots of intact maize seedlings Pestic. biochem. physiol.43: 212-222 Kasai and Bayer. 1991. J. Pestic. Sci. 16:171-177. 6. Hess, F.D. 1999. Surfactants and additives. Proceedings of the California Weed Science Society 51: 156-172.

7. Sterling T.M. 1994. Mechanisms of herbicide absorption across plant membranes and accumulation in plant cells, Weed Sci. 42:263-276. CITOCROMO P450 MONOXIGENASA 1. Barrett, M. 2000. The role of cytochrome P450 enzymes in herbicide metabolism. Pp. 24-37 In A. H. Cobb and R.C. Kirkwood Herbicides and their Mechanisms of Action. Boca Raton, FL: CRC Press. 2. Davies, J. and J. Caseley. 1999. Herbicide safeners: a review. Pestic. Sci. 55:10431058. 3. Devine, M., S.O,. Duke, and C. Fedtke.1993. Physiology of Herbicide Action. Englewood Cliffs, New Jersey: Prentice Hall. Pp. 95-112. 4. Mougin, C.P., M.-F. Corio-Costet, and D. Werk-Reichhart. 2000. Plant and fungal cytochrome P-450s: their role in pesticide transformation. Pp. 166-181 In J.C. Hall, R.E. Hoagland, and R.M. Zablonowicz, Pesticide Biotransformation in Plants and Microorganisms, Similarities and Divergences. Washington, DC: American Chemical Society. 5. Preston, C. and S. Powles. 1997. Light-dependent enhanced metabolism of chlorotoluron in a substituted urea herbicide-resistant biotypr of Lolium rigidum. Gaud. Planta 201: 202-208 6. Preston et al. 1996. Multiple resistance to dissimilar herbicide chemistries in a biotype of Lolium rigidum due to enhanced activity of several herbicide degrading enzymes. Pestic. Biochem. Physiol. 54:123-134. FASE II DEL METABOLISMO DE HERBICIDA EN LA PLANTA 1. Edwards, R. and D.P. Dixon. 2000. The role of glutathione transferases in herbicide metabolism. Pp. 38-71 In A. H. Cobb and R.C. Kirkwood Herbicides and their Mechanisms of Action. Boca Raton, FL: CRC Press. 2. Davies, J. and J. Caseley. 1999. Herbicide safeners: a review. Pestic. Sci. 55:10431058. 3. Devine, M., S.O,. Duke, and C. Fedtke.1993. Physiology of Herbicide Action. Englewood Cliffs, New Jersey: Prentice Hall. Pp. 95-112. 4. Hatzios, K.K. 2000. Functions and regulations of plant glutathione S-transferases. Pp. 218-239 In J.C. Hall, R.E. Hoagland, and R.M. Zablonowicz, Pesticide Biotransformation in Plants and Microorganisms, Similarities and Divergences. Washington, DC: American Chemical Society. 5. Hatzios, K.K. and N. Burgos. 2004. Metabolism-based herbicide resistance: regulation by safeners. Weed Sci. 52:454-467 6. Romano, M.L., Gerald R. Stephenson, Abraham Tal, and J. Christopher Hall. 1993. The effect of monooxygenase and glutathione S-transferase inhibitors on the metabolism of diclofop-methyl and fenoxaprop-ethyl in barley and wheat. Pestic. Biochem. Physiol. 46:181-189. INHIBIDORES DE LA ALS

1. Ahrens, W.H. and W.R. Panaram. 1997. Weed Sci. 45:648-653. 2. Brown, H.M. 1990. Pestic. Sci. 29: 263-281. 3. Davis, J and JC Caseley. 1999. Pesticide Science 55:1043-1058 4. Devine, M., S.O,. Duke, and C. Fedtke.1993. Physiology of Herbicide Action. Englewood Cliffs, New Jersey: Prentice Hall. 5. Duggleby RG and SS Pang, 2000. J. Biochem and Mol. Bio. 33:1-36 6. Dyer, WE, PW Chee and PK Fay, 1993. Weed Sci. 41:18-22 7. Geiger, DR and HD Bestman. 1990. Weed Science 38:324-329. 8. Hall, L.M. and M.D. Devine. 1993. Pestic. Biochem. Physiol. 45:81-90 9. Hart, S.E. and D. Penner. 1993. Weed Sci. 41: 28-33. 10. Herbicide Handbook. 2002. WSSA, 8th ed. 11. Kreuz,K. and R. Fonne-Pfister. 1992. Pestic. Biochem. Physiol. 43:232-240 12. McCourt JA, SS Pang, JK Scoot, LW Guddat and RG Duggleby, 2006. PNAS 103(3):569573. 13. Pang SS, RG Duggleby and LW Guddat, 2002. J. Mol. Bio. 317:249-262 14. Saari, LL, JC Cotterman and DC Thill, Resistance to ALS inhibiting herbicides, in Herbicide resistance in plants, 1994, CRC Press. pp.83-139 15. Shaner, DL and BK Singh, AHAS inhibitors, in Herbicide activity: Toxic., Biochem. and Mol Bio., pp. 69-110 16. Tranel PJ and TR Wright, 2002. Weed Science 50:700-712

INHIBIDORES DE SNTESIS LIPDICA 1. Brommer, C.L. et al. 2000. Antagonism of BAS 625 by selected broadleaf herbicides and the role of ethanol. Weed Sci. 48:181-187. 2. Boger, P. et al. 2000. Towards the primary target of chloroacetamides-new finding pave the way. Pest Manag. Sci. 56:497-508. 3. Burton, J.D. 1997. Acetyl-Coenzyme A Carboxylase inhibitors. Pgs. 187-205 In R.M. Roe et al. (eds) Herbicide Activity: Toxicology, Biochemistry, and Molecular Biology. Amsterdam, Netherlands: IOS Press. 4. Christopher, J.T. and J.A.M Holtum. 2000. Dicotyledon lacking the multisubunit form of the herbicide-target enzyme acetyl coenzyme A carboxylase may be restricted to the family Geraniaceae. Aust. J. Plant Physio. 27: 845-850. 5. Delye, C. 2005. Weed resistance to acetyl coenzyme A carboxylase inhibitors: an update. Weed Sci. 53: 728-746. 6. Delye, C. et al. 2005. Molecular bases for sensitivity to Acetyl coenzyme A caroxylase inhibitors in black grass. Plant Physiol. 137:794-806. 7. Devine, M., S.O. Duke, and C. Fedtke. 1993. Physiology of Herbicidal Action. Englewood Cliffs, New Jersey: Prentice Hall. 8. DiTomaso, J.M. 1994. Evidence against a direct membrane effect in the mechanism of action of graminicides. Weed Sci. 42: 302-309. 9. Gonwald, J.W. 1991. Lipid Biosynthesis Inhibitors. Weed Sci. 39:435-449.

10. Incledon, B.J., and J.C. Hall. 1997. Acetyl Coenzyme A Carboxylase: Quatanary structure and inhibition by graminicidal herbicides. Pest. Biochem. and Physiol. 57: 255271. 11. LeTouze, A., and .J Gasquez. 1999. A pollen test to detect ACCase target-site resistance within Alopecurus myosuioides population. Weed Res. 40:151-162.` 12. Jachetta, J. 2005. Lipid Biosynthesis Inhibiting Herbicides. Presentation from Weed School. UC Davis. 13. Moss, S.R . et al. 2003. Characterization to target-site resistance to ACCasie inhibiting herbicides in the weed Alopecurus mysosuroides (black-grass.) Pest Manag. Sci. 59:190-201. 14. Romano, M.L. et al. 1993. The effect of monooxygenase and glutathione S-Transferase inhibitors on the metabolism of diclofop-methyl and fenoxaprop-ethyl in barely and wheat. Pest. Biochem. and Physiol. 46:181-189. 15. Ross, M.A., and C.A. Lembi. 1999. Applied Weed Science. 2nd. Ed. Upper Saddle River: Prentice Hall. 16. Sasaki, Y. and Y. Nagano. 2004. Plant acetyl CoA carboxylase: structure, biosynthesis, regulation, and gene manipulation for plant breeding. Biosci. Biotechnol. and Biochem. 68(6):1175-1184. 17. Gressel, J. 2002. Molecular Biology of Weed Control. London: Taylor & Francis. Chapter 4. HERBICIDAS QUE GENERAN FORMAS TXICAS DE OXGENO 1. Taiz, L. & E. Zeiger. 2002. Plant Physiology-Third Ed. Sunderland, Massachussetts: Sinauer. Chapter 7. 2. Devine, M., S.O. Duke, and C. Fedtke. 1993. Physiology of Herbicidal Action. Englewood Cliffs, New Jersey: Prentice Hall. Chapter 7, 8 and 9 3. Hess, F. D. 2000. Light-dependent herbicides: an overview. Weed Sci. 48:160-170 4. Chul, J-C. 2001. Comparative absorption, translocation, and metabolism fo foliarapplied oxyfluorfen in wheat and barley. Pestic. Biochem. Physiol. 70:118-125. 5. Pallett, K.E. 2000. The mode of action of isoxaflutole: a case study of an emerging target site. Pages 215-238 in A.H. Cobb and R.C. Kirkwood (eds.) herbicides and their mechanisms of Action. Sheffield, England: CRC Press. 6. Szigeti, Z. & E. Lehoczki. 2003. A review of physiological and biochemical aspects of resistance to atrazine and paraquat in Hungarian weeds. Pest Manag. Sci. 59:451-458. 7. Sandmann, G. and P. Bger. 1997. Phytoene desaturase as a target for bleaching herbicides. Pages 1-10 in Roe et al. (eds.) Herbicide Activity: toxicology, Biochemistry and Molecular Biology. Amsterdam, Netherlands: IOS Press. 8. ElNaggar, S.F. et al. 1992. Metabolism of clomazone herbicide in soybean. J. Agric. Food Chem. 40:880-883. 9. Mueller, C. et al. 2000. Properties and inhibition of the first two enzymes of the nonmevalonate pathway of isoprenoid biosynthesis. Biochemical society Transactions. 28(2): 792-793. 10. Culpepper, A.S. et al. 2001. Effect of insecticides on clomazone absorption, translocation, and metabolism on cotton. Weed Sci. 49:613-616. 11. Zeidler et al. 2000. Biochem. Soc. Trans. 28: 796-798

12. Ferhatoglu et al. 2005. Pesticide Biochem. Physiol. 81:59-70 13. Ferhatoglu & M. Barrett. 2006. Pesticide Biochem. Physiol. 85: 7-14. INHIBIDORES MITTICOS 1. Devine, M., S.O. Duke, and C. Fetdke. 1993. Physiology of Herbicide Action. Englewood Cliffs, , New Jersey: Prentice hall. Chapter 10. 2. Hoffman, J.C. and K.C. Vaughn. 1994. Mitotic disrupter herbicides act by a single mechanism but vary in efficacy. Protoplasma 179:16-25. 3. Molin, W.T. and R.,A. Khan. 1997. Mitotic disrupter herbicides: Recent advances and opportunities. Pages 143-158 in R.M. Roe et al. (eds.) Herbicide Activity: Toxicology, Biochemistgry and Molecular Biology. Amsterdam, Netherlands: IOS Press. 4. Hess, F.D. 1989.Herbicide interference with cell division in plants. Pages 85103. In: Target Sites of Herbicide Action. Boger, P. and G. Sandmann, eds. Boca Raton, FL: CRC Press Inc. 5. Ellis, J., R. Taylor and P. Hussey, 1994. Molecular modeling indicates that two chemically distinct classes of anti-mitotic herbicides bind to the same receptor site(s). Plant Physiol. 105:518. 6. Desai, A., and T. Mitchision, 1997. Microtubule polymerization dynamics. Ann. Rev. Cell and Develop. Biol. 13:83117. 7. Taiz, L. and D. Zeiger. 1998. Plant Physiology. Sunderland, Massachussets: Sinauer Assoc., Inc., Publ.

HERBICIDAS AUXNICOS 1. M.D. Devine, S.O. Duke and C. Fedtke. 1993 Physiology of Herbicide Acion. PTR Prentice Hall, Englewood Cliffs, New Yersey, 07632. Chapter 14 2. Sterling, T.M. and J. Christopher Hall. 1997. Mechanism of action of natural auxins and the auxinic herbicides. Pages 111-141 In R.M. Roe et al. (eds.) Herbicide Activity: Toxicology, Biochemistry and Molecular Biology. Amsterdam: IOS Press. 3. Abdallah, I., A.J. Fischer, C.L Elmore, M. E.Saltveit, and M. Zaki. 2006. Mechanism of Resistance to Quinclorac in Smooth Crabgrass (Digitaria ischaemum). Pesticide Biochemistry and Physiology 84:38-48.. 4. Taiz, L. and Zeiger, E. (eds.). 2002. Plant Physiology, Sinauer Assocs., Sunderland, MA. Chapter on auxins. 5. Grossman, K. 2000. The mode of action of quinclorac: a case-study of a new auxi-type herbicide. Pp 181-214 A.C. Cobb and R.C. Kirkwood (eds.). Herbicides and Their Mode of Action.CRC Press: Glasgow, UK. 6. Grossman, K. and J. Kwiatkowski. 2000. The mechanism of quinclorac selectivity in grasses. Pestic. Biochem. Physiol. 66:83-91. 7. Zheng, H., J. C. Hall. 2001. Understanding auxinic herbicide resistance in wild mustard: physiological biochemical, and molecular genetics approaches. Weed Sci. 49:276-281.

GLIFOSATO 1. Geiger, D.R. et al. 1986. Glyphosate inhibits photosynthesis and allocation of carbon to starch in sugar beet leaves. Plant Physiol. 82:468-472. 2. Geiger, D.R. and M. A. Fuks. 2002. Inhibitors of aromatic amino acid biosynthesis (Glyphosate). Pages 59-85 In P. Boger et al. (eds.) Herbicide Classes in Development Mode of Action, Targets, Genetic Engineering, Chemistry. Berlin: Springer. 3. Siehl, d.L. 1997. Inhibitors of EPSP synthase, glutamine synthetase and histidine synthesis. Pages 37-67 In R.M. Roe et al. (eds.) Herbicide Activity: toxicology, biochemistry and Molecular Biology. Amsterdam: IOS Press. 4. Devine, M. and c. Preston. 2000. The molecular basis of herbicide resistance. Pages 8586 In A.H. cobb and R.C. Kirkwood (eds.) Herbicides and their Mechanisms of Action. Sheffield, UK: CRC Press. 5. Wayakabayashi, K. and P. Boger. 2004. Phytotoxic sites of action for molecular design of modern herbicides (Part 2): amino acid, lipid and cell wall biosynthesis, and other targets for future herbicides. Weed biol. Manage. 4:59-70. 6. T. T. Lee Release of Lateral Buds from Apical Dominance by Glyphosate in Soybean and Pea Seedlings J Plant Growth Regul (1984) 3:227-235 7. JAMES H. WESTWOOD and DAVID D. BIESBOER The influence of glyphosate on endogenous levels of free IAA and phenolic compounds in leafy spurge Reprinted from: 1985 Leafy Spurge Symposium. Bozeman, MT. July 17-18, 1985. pp. 5-13 8. Marie-Hlne Denis1 and Serge Delrot1 Carrier-mediated uptake of glyphosate in broad bean (Vicia faba) via a phosphate transporter Physiologia Plantarum Volume 87 Page 569 RESISTENCIA DE MALEZAS A HERBICIDA 1. Cousens, R. and M. Mortimer. 1995. Dynamics of weed populations. Cambridge, UK: Cambridge University Press. Pp. 243-282. 2. Di Tomaso, J.M. 2001. Herbicide resistance. In Weed Science School, september 2628, 2001, University of California, Davis. 3. Fischer, A.J., Ateh, C. M., Bayer, D. E., Hill, J. E. 2000. Herbicide-resistant Echinochloa oryzoides and E. phyllopogon in California Oryza sativa fields. Weed Science 48: 225230. 4. Gressel, J. and L.A. Segel. 1978. The paucity of plants evolving genetic resistance to herbicides: possible reasons and implications. J. Theor. Biol. 75:349-371. 5. Gressel, J., Molecular Biology of Weed Control, pp 115-117 and 200-201, London and New York, Taylor and Francis, 2002. 6. Guttieri, M.J., C.V. Eberlein, D. Thill. 1995. Diverse mutations in the acetolactate synthase gene confer chlorsulfuron resistance in kochia (Kochia scoparia) biotypes. Weed Sci. 43: 175-178. 7. Jasieniuk, M., A.L. Brl-Babel, and I.M. Morrison. 1996. The evolution and genetics of herbicide resistance in weeds. Weed Sci. 44:176-193. 8. Maxwell, B. D., M.L. Roush, and S.R. Radosevich. 1990. Predicting the evolution and dynamics of herbicide resistance in weed populations. Weed Technol. 4:2-13.

9. McCourt, J.A., S.S. Pang, J.King-Scott, L.W. Guddat, and R.G. Duggleby. 2006. Herbicide-binding sites frevealed in the structure of plant acetohydroxyacid synthase. PNAS 103:569-573 10. Osuna, M.D., F. Vidotto, A.J. Fischer, D.E. Bayer, R. De Prado, and A. Ferrero. 2002. Cross-resistance to bispyribac-sodium and bensulfuron-methyl in Echinochloa phyllopogon and Cyperus difformis. Pestic. Biochem. Physiol. 73:9-17. 11. Tsuji, R., A.J. Fischer, M. Yoshino, A. Roel, J.E. Hill and Y. Yamasue. 2003. Herbicideresistant late watergrass (Echinochloa phyllopogon): similarity in morphological and amplified fragment length polymorphism traits, Weed Sci. 51 (2003) 740-747. MEZCLAS DE HERBICIDAS 1. Colby, S.R. 1967. Calculating synergistic and antagonistic responses of herbicide mixtures. Weeds 15:20-22. 2. Brain, P., and R. Cousens. 1989. An equation to describe dose responses where there is stimulation of growth at low doses. Weed Research 29:93-96. 3. Fischer, A.J., D.P. Cheetham, and R. De Prado. 2004. Enhanced effect of thiobencarb on bispyribac-sodium control of E. phyllopogon (Stapf) Koss in California rice (Oryza sativa L.). Weed Biology and Management 4:206-212. 4. Gressel, J. 1990. Synergizing herbicides. Rev. Weed Sci. 5:49-82. 5. Gressel, J. 2000. Molecular biology of weed control. Transgenic Research 9:355-382. 6. Limpel, L.E., P.H. Schuldt, and D. Lamont. 1962. Weed control by dimethyl tetrachloroterephthalate alone and in certain combinations. Proc. Northeastern Weed Control conf. 16:48-53. 7. Schabenberger O & Birch J (2001) Statistical dose-response models with hormetic effects. Human and Ecological Risk Assessment, 7, 891-908. 8. Streibig J, Rudemo M & Jensen J (1993) Dose-response curves and statistical models. In: Herbicide Bioassays. pp. 30-55: CRC Press, Boca Raton, FL.