Mechanisms of Oryza sativa (Poaceae) resistance to Tagosodes
Contents
1. B 5 Poy l A 5 s s 5 3 E h s a E 5 a i a fo e 3 n 3 S wo 3 br u 2 Y E o s gt oO gt 2 x ES A 7 2 8 E 30 B i 2 lt 25 es 3 ER Eg g ge 15 23 10 os i H l i m 2 g s 8 E amp 2 S 4 E 8 g g 2 E R amp 3 z 2 z lt s e s E 3 s x n 5 lt NA S 3 2 ne 2 2 gt 2 ES g 3 5 x 8 E 2 e a 35 C a lt 30 Nc No DE 25 Eg A 204 ES E 15 5 Oe 10 2 ml H 0 O o o oO o E E rs 5 2 2 2 El 2 E A 2 2 S 3 g gt a lt T S E 3s i nt 2 a 3 2 ES g i z a a a 2 Rice genotypes Fig 1 Antixenosis Average of settling nymphs of Tagosodes orizicolus after 24 A 48 B and 72 C hours after infestation Bars with same letters do not differ significantly a 0 05 cultivars Kruskal Wallis p 0 0024 being Bluebonnet 50 Araure 4 and D sati va which showed the highest susceptibil ity level to direct damage On the other hand PNOOAO16C Centauro PFD 2 1 1 2 gt 293 M 6 2 2 1 and FD0241 M 17 6 1 1 1 1 showed intermediate resistance level while Makalioka FD0241 M 18 3 1 2 M 1 Rev Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 60 1 105 117 March 2012 Fedearroz 50 and Venezuela 21 exhibited th2. Mechanism of rice genotypes resistance to T orizicolus From the agricultural perspec tive a resistant cultivar is defined when it yields is better under the pressure of an insect plague compared with a susceptible one Niks amp Lindhout 1999 Thus the resistance of a plant to a plague is a relative term based on the comparison with a susceptible plant Niks amp Linhout 1999 Baddi amp Garza 2007 In this research a thorough comparison of the differ ent groups was made and it was possible to see that all cultivars expressed some kind of resistance mechanism differing in type and expression level For example Makalioka showed higher levels of resistance to direct damage produced by sogata this resistance was determined by the antixenosis mechanism which increased as the exposition time to insect was longer besides survival and oviposition antibiosis were high coinciding with other reports Pardey et al 1996b Calvert et al 2004 The cultivars Fedearroz 50 and Venezu ela 21 are susceptible to direct damage even though they have high antixenosis and medium antibiosis for survival and oviposition under greenhouse conditions D sativa has medium antixenosis and medium high antibiosis for survival and oviposition however it becomes susceptible to direct damage at certain insect population densities Centauro is a moderately resistant cul tivar to direct damage The resistance is pro vided by strong anti
3. F E pra s 5 lt Y S lt 5 T N T Ss a E 2 eo Pol a z 00 D o mM a 3 _ e n gt a z A x 3 S E a a a Rice genotypes Fig 2 Antibiosis Evaluation of genotype effect over survival of Tagosodes orizicolus Bars with same letters do not differ significantly a 0 05 Rev Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 60 1 105 117 March 2012 111 m Free choice No choice s gg Oo gs S 8 71 S S S OF E mE uv 5 a 4 E RH 1 l E 34R R R R zo ES uv 14 Q g 0 Z R A 8 Y Za 3 sg 8 2 N 2 2 8 a 5 g 2 NN E U E lt eE 4 lt ES v T s vr g gg s lt 2 6 3 2 Zz T 2 2 2 68 amp 2 A Sa a lt Le w Q a FD0241 M 17 6 1 1 1 1 Rice genotypes Fig 3 Average of direct damage produced by Tagosodes orizicolus on rice cultivars and elite lines in row free choice and no choice tests showed lower level of antibiosis for oviposi tion than Araure 4 and Bluebonnet 50 despite it exhibited moderate level of antibiosis for survival Tolerance in greenhouse conditions The FPLI showed statistical differences between treatments p 0 043 seeing that some geno types lost a significant amount of biomass as a result of insect infection Differences related to the FPLI and direct damage in each treat ment was observed through mean test Table 1 Considering the FPLI and direct damage Centauro
4. was the most tolerant cultivar to T orizicolus because it had one of the lowest FPLI with a considerable level of direct dam age Several genotypes showed low FPLI but associated with a low level of direct damage The line FD0241 M 17 6 1 1 1 1 showed a lower FPLI than susceptible Bluebonnet 50 with direct damage mean of 4 50 which could be indication of tolerance for that genotype DISCUSSION Antixenosis using free choice test This method proposed by Heinrichs et al 1985 to detect antixenosis in rice for Nilaparvata 112 lugens Stal Delphacidae Nephotettix vires cens Distant Cicadellidae Sogatella furcifera Horvath Delphacidae y Recilia dorsalis Mot shulsky Cicadellidae was effective to study feeding preference of T orizicolus under green house conditions There are few references related to free choice feeding using circle arrangements in rice exposed to T orizicolus however the num ber of settling insect expressed as percentage at 24 36 and 72h were similar to those reported by Calvert et al 2004 where Fedearroz 50 and Makalioka were consistently less pre ferred by sogata The results from free choice feeding in the row arrangement were similar to previous reports where Makalioka and Bluebonnet 50 were more resistant and susceptible respec tively Triana et al 1994 High susceptibility was shown by Araure 4 using this method under greenhouse condition
5. identified three mecha nisms of plant resistance to insects antibiosis tolerance and non preference Subsequently other researchers recognized the behavior of insects associated with non preference the category called antixenosis has been widely accepted Panda amp Khush 1995 Each one of these three main categories of plant resistance may act by different mechanisms The genetic characteristics of the plant can be modified by conventional breeding or by the use of biotechnology techniques which allow the flow of genes between selected plants Badii amp Garza 2007 Rice breeding programs have focused into obtaining new cultivars with increased resistance to direct and indirect dam age produced by the insect However few studies has been made on the mechanisms of resistance in rice Pardey et al 1996b Alvarez et al 2000 Pardey et al 2000 Calvert et al 2004 Calvert et al 2005 Many cultivars used as source of resistance to sogata in breeding programs have not been characterized for their resistance type On the other hand it have been used resistance sources that present the antibiosis mechanism which is undesirable because sogata is able to overcome resistant in few generations Pantoja et al 1993 Pantoja et al 1997 Improvement plants resistance to RHBV and its vector is considered of paramount importance in rice breeding programs in Ven ezuela because sogata populations increase considerably during the rice
6. Rice Research Institute Los Ba os Laguna Philippines Hern ndez M T Quesada C Mu oz amp A M Espinoza 2004 Genetic diversity of Costa Rican populations of the rice planthopper Tagosodes orizicolus Homopte ra Delphacidae Rev Biol Trop 52 795 806 Rev Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 60 1 105 117 March 2012 115 Infostat 2004 Infostat Versi n 2004 Universidad Nacio nal de C rdoba C rdoba Argentina IRRI 2002 Reference Guide Standard Evaluation Sys tem for Rice International Rice Research Institute Consultado 2 mayo 2008 http www knowledge bank irri org ses Labr n N A Ebert I P rez Almeida C Astorga G Rivas amp L Calvert 2009 Evaluaci n de la resisten cia de cultivares de arroz venezolanos ante el virus de la hoja blanca y su asociaci n con marcadores microsat lites Fitopatol Venez 22 13 18 Malaguti G 1956 La hoja blanca extra a enfermedad del arroz en Venezuela Agronom a Trop 6 141 145 Mart nez C 2005 Hacia la comprensi n de la base gen tica de la resistencia al complejo del virus de la hoja blanca y Tagosodes orizicolus en arroz Res menes de la XLV Reuni n Anual de la APS San Jos Costa Rica Meneses C R Y A Guti rrez R A Garc a P G Antigua S J G mez F Correa amp L Calvert 2001 Gu a para el trabajo de campo en el manejo integrado de plagas del arroz Colombia FLAR Painter R H
7. caused by this pest Vivas et al 2002 reducing over 40 of rice production in Venezuela since its appearance Malaguti 1956 Rev Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 60 1 105 117 March 2012 The males of T orizicolus sogata are about 2mm long dark brown or black and the females are between 3 33 and 3 35mm long amber lighter than males the back of the chest to the keel is paler and this color extends to the head The eggs are transparent and are between 0 50 and 0 70mm long The nymphs are whit ish in color and their size varies between 0 65 0 60mm long x 0 20 0 30mm wide As they grow the sharpness of the lateral lines of their body increases Meneses et al 2001 The females can lay up to 350 eggs in the spongy parenchyma into the leaf midrib The hatching 105 can take between four and eight days but it can vary depending on temperature Nymphs go through five stages whose duration can vary between 15 and 20 days Finally adults have a life span ranging from 14 to 36 days Pantoja et al 1997 The symptoms of direct damage are accel erated leaf senescence and leaf necrosis which begin from the apex and edges to the basal part of the leaf reducing growth and eventually causing the death of the plants lvarez et al 2000 Resistance to direct damage caused by sogata is controlled by the combined action of a major gene with some modifier genes Pardey et al 2000 Romero 2005 Painter 1951
8. 1951 Insect resistance en crop plants Mac Millan New York USA Panda N amp G S Khush 1995 Host plant resistance to insects CAB International amp International Rice Research Institute Manila Philippines Pantoja A A Fischer F Correa Victoria L R Sanint amp A Ram rez 1997 MIP en Arroz Manejo integrado de plagas artr podos enfermedades y malezas Centro Internacional de Agricultura Tropical Cali Colombia Pantoja A R Zeigler amp F Correa 1993 Screening rice germoplams for Tagosodes orizicolus resistance and detecting the percentage of vector p 53 55 In Pro ceeding of monitoring tour and workshop on integrate management of rice in the Caribbean CIAT eds Cali Colombia Pardey C D Baena amp F Cuevas 19964 Herencia de la resistencia al da o mec nico causado por Tagosodes orizicolus Muir Homoptera Delphacidae en arroz Oryza sativa Acta Agron 46 9 14 Pardey C F Cuevas C Martinez amp A Pantoja 2000 Herencia de la resistencia al da o mec nico por Tago sodes orizicolus Muir Homoptera Delphacidae en arroz J Agri Univ Puerto Rico 84 47 52 Pardey C F Cuevas D Baena amp C Mart nez 1996b Caracterizaci n de la resistencia al da o mec nico de Tagosodes orizicolus Muir Homoptera Delphaci dae en doce cultivares de arroz Oryza sativa Rev Col Entomol 22 37 43 Romero L E 2005 Aproximaci n hacia la identificaci n de marcado
9. 5 febrero 2007 http www redpav fpolar info ve danac volumen6 art 1 index html Vivas L E L Lugo M Acevedo amp S Clavijo 2002 Determinaci n de la preferencia de Tagosodes orizi colus MUIR 1926 Homoptera Delphacidae sobre variedades de arroz en Calabozo estado Gu rico Venezuela Investigaci n Agr cola 7 1 Consultado 28 febrero 2007 http www redpav fpolar info ve danac viewarticle php id 33 Vivas L E D Astudillo amp L Campo 2007 Evaluaci n del insecticida Etofenprox 10 9 para el control del insecto sogata en el cultivo del arroz en Calabozo Estado Gu rico Venezuela Agronom a Trop 57 287 297 116 Rev Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 60 1 105 117 March 2012 Vivas L E D Astudillo amp L Campo 2009 Evaluaci n del insecticida Thiamethoxan 25 para el manejo del insecto sogata en el cultivo del arroz en Cala bozo Estado Gu rico Venezuela Agronom a Trop 59 89 98 Weber G amp O Parada 1994 Development of an Inte grated Pest Management System for rice in Latin America p 733 748 In E A Heinrichs ed Bio logy and management of rice insects Wiley Eastern Limited New Delhi India Zeigler R S A Pantoja M C Duque amp G Weber 1994 Characteristics of resistance in rice to rice hoja blanca virus VHB and its vector Ann Appl Biol 124 429 440 Rev Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 60 1 105 117 Ma
10. Araure 4 and Bluebonnet 50 did not show any deleterious effects on sogata the other genotypes showed intermediate and high level of antibiosis Fig 2 Comparison of direct damage between free choice in row and no choice test There were no significant differences in direct dam age p 0 1631 between mean treatments in no choice test However the mean value over all the genotypes was used to compare both feeding methods Fig 3 The cultivars Ven ezuela 21 Fedearroz 50 and FD0241 M 18 3 1 2 M 1 showed typical reactions of high antixenosis where the genotypes showed resis tance in free choice and were susceptible or intermediate in no choice conditions Maka lioka Centauro FDO241 M 17 6 1 1 1 1 and PFD 2 1 1 2 gt 293 M 6 2 2 1 were the cultivars that showed more stable resistance to sogata across both methods Antibiosis using no choice test for ovi position In this trial statistical differences p 0 0006 between cultivars were detected Six different groups were obtained with the Duncan s test Table 1 where Makalioka Centauro and PNOOAO16C exhibited higher antibiotic effect and FDO241 M 17 6 1 1 1 1 had lowest effect on oviposition of the insects The genotype FDO241 M 17 6 1 1 1 1 905 C 80 BC 70 BC 6 ae AB Lo AB 2 504 AB AB AB ho Y 40 Y 304 i A un 20 104 i 01 _ 2 2 zs 8 ZS 3 3 u q N S g
11. MOL UT 9DIOYI 991H I S Y I Y I a a a a Y oseulep JO AT Voge a Ogl V 00 Voge V 00 av 00 f V 00 V 00 V 00 V 00 V 00 cn xP IPO IYOMO UT DDIOYI 391H SISOUIXNUY I T I T 9 Z1T W T c0da Os 19uuoq n q I IN C I 91 IN IFC0dH I c 9 IW 6 lt AINV dHd IZ Bjanzous y amery OS ZOLIB9po Panes d Ao e ela omego 3910Y 00Nd sodAjousy Rev Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 60 1 105 117 March 2012 110 seven resistant varieties three with interme diate reaction and one susceptible could be differentiated whereas in the row feeding experiment a greater number of susceptible genotypes were observed PNOOA016C and Centauro changed from resistant in the circle feeding experiment to susceptible in the row feeding exposure The most drastic effect was observed on D Sativa that changed damage level from resistant to susceptible A simi lar result was observed in Araure 4 which changed from intermediate level observed in the circle method to susceptible in row method Antibiosis using no choice test for sur vival Significant differences p 0 0044 on insects mortality were detected between rice genotypes using no choice test for survival The Duncan mean test showed four homo geneous groups for survival effect on sogata Makalioka exhibited the highest antibiosis level where 83 33 of adult insects died while
12. Mechanisms of Oryza sativa Poaceae resistance to Tagosodes orizicolus Homoptera Delphacidae under greenhouse condition in Venezuela Alex Gonz lez Natalia Labr n Rosa M lvarez Yorman Jayaro Carlos Gamboa Edicta Reyes amp Venancio Barrientos L Fundaci n para la Investigaci n Agr cola Danac Carretera San Javier v a Guarataro Finca Naranjal Apdo 182 Felipe Venezuela alex gonzalez danac org ve natalia labrin danac org ve yorman jayaro danac org ve venancio barrientos danac org ve Instituto Nacional de Investigaciones Agropecuarias INIA Araure Venezuela rmalvarez inia gob ve ereyesQinia gob ve Received 22 X1 2010 Corrected 03 VI 2011 Accepted 07 VH 2011 Abstract Tagosodes orizicolus is one of the main plagues of rice in tropical America causing two types of damages the direct one feeding and oviposition effect and an indirect one by the transmission of the Rice hoja blanca virus During 2006 2007 we carried out research under greenhouse conditions at Fundaci n Danac Venezuela in order to determine the mechanisms of antixenosis antibiosis and tolerance to T orizico lus which could be acting in commercial varieties and advanced lines of the rice genetic breeding programs of INIA and Fundaci n Danac The method of free feeding was used for the antixenosis evaluation whereas the method of forced feeding was used for antibiosis evaluation effect on survival and oviposition Additionally we use
13. a nisms identifying itself as resistant progenitor 114 Rev Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 60 1 105 117 March 2012 to T orizicolus to be use in the breeding pro grams in Venezuela ACKNOWLEDGMENTS Authors want to thank to Fondo Nacional de Ciencia Tecnolog a e Innovaci n FONAC IT of Venezuela for partial funding given to Project N 2000001658 entitled Rice Germo plasm Evaluation resistance to direct damage produce by sogata Tagosodes orizicolus Muir and hoja blanca virus whose partial results are presented in this paper RESUMEN Tagosodes orizicolus es una de las principales plagas del cultivo del arroz en Am rica tropical causando dos tipos de da os a la planta el directo por efecto de la ali mentaci n y oviposici n el indirecto por la transmisi n del virus de la hoja blanca del arroz Durante el per odo 2006 2007 se llev a cabo una investigaci n bajo condi ciones de umbr culo en la Fundaci n Danac Venezuela con el objeto de determinar los mecanismos de resistencia de antixenosis antibiosis y tolerancia a T orizicolus que pudieran estar actuando en l neas avanzadas y cultivares del programa de mejoramiento gen tico de arroz de las instituciones INIA y Fundaci n Danac Para la evaluaci n de antixenosis se utiliz el m todo de libre elecci n para antibiosis efecto sobre sobrevivencia y oviposici n de los insectos el de alimentaci n forzada adicionalmente s
14. al mortality of insects Alvarez et al 2000 Direct damage in plants using no choice test for survival The conditions and design method in this assay were the same used in the latter experiment The infestation was made Rev Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 60 1 105 117 March 2012 107 only with five pairs of young adults insects 21 25 days after hatching per plot those plants were not shacked out Direct damage in each plant was evaluated 30 days after infestation to determine the genotype reaction under forced feeding conditions lvarez et al 2000 Phe notypic evaluation was done using the Standard System Evaluation for Rice IRRI 2002 Comparison of direct damage between free choice in row and no choice test The mean of the direct damage of genotypes obtained in the free choice in row test and no choice test for survival were compared Antibiosis using no choice test for ovi position For each rice genotype five seeds per plot were planted in pots of 9 50cm x 11 50cm under a complete randomized design with four replications One plant per plot was left eight days after planting and fertilized with urea 29 days after planting plants were covered with micro cages made with 2L clear plastic bottles and then a pair of young adult insects was release Ten and twenty days after infesta tion replacement of genotype was made and the number of eggs was counted in each plant dissecting the tis
15. cm between rows were planted A completely randomized design with four replications was used Five plants were left in the row and fertilized with urea eight days after planting 25 days after planting trays were placed into an entomological cage 80cm height x 100cm length x 50cm deep and plants were infested with five nymphs per plants at the fourth and fifth instar Plants were shacked out every day until 30 days after infestation The evaluation of direct damage was made when Bluebonnet 50 reached the highest in the range according to the Standard System Evalu ation for Rice IRRI 2002 Antibiosis using no choice test for sur vival Ten seeds of each genotype were planted in pots of 9 50cm x 11 50cm under a com pletely randomized design with three replica tions Plants were thinned to five per pot and fertilized with urea eight days after planting 15 days after planting plants were covered with micro cages made with 2L clear plastic bottles then the plants were infested with 10 pairs of young adult insects 21 to 25 days after hatching per pot the number of dead insects per plot was counted daily until all of them died Dead insect data obtained were trans formed to percentage of mortality The analysis was made with the percentage of mortality at eight days after infestation when the mortality of insect on the susceptible cultivar Bluebon net 50 reached 20 it was made to avoid the antibiosis effect from the natur
16. d the indirect method based on biomass depression to estimate the tolerance Some of the evaluated traits included grade of damage number of insects settling on rice plants percentage of sogata mortality at the mature state number of eggs in the leaf midrib and an index of tolerance The results showed that rice geno types possess different combinations of resistance mechanisms as well as different grades of reactions The susceptible control Bluebonnet 50 was consistently susceptible across experiments and the resistant control Makalioka had high antixenosis and high antibiosis based on survival and oviposition The rest of the geno types presented lower or higher degrees of antixenosis and antibiosis for survival and oviposition The genotype FDO0241 M 17 6 1 1 1 1 was identified with possible tolerance to the direct damage of sogata Rev Biol Trop 60 1 105 117 Epub 2012 March 01 Key words rice sogata Rice hoja blanca virus RHBV antixenosis antibiosis tolerance San Rice in tropical America is affected by the planthopper Tagosodes orizicolus Muir Common name sogata causing direct dam age when the nymphs and adults feed from the mesophyll and phloem sap at different stages of plant development or when the adults oviposit on the plant leaves Zeigler et al 1994 An indirect damage can also be caused by sogata as vector of the Rice hoja blanca virus RHBV which is one the most important disease
17. e highest resistance level to direct damage caused by T orizicolus Table 1 The circle and row feeding experiments showed contrasting results regarding genotype resistance and susceptibility to mechanical damage In the circle feeding experiment 109 10 0 gt d s ou jjJip DANBOLIUSIS SOJLOLPUL 199 JUSIJI S861 12 suoutut H x pul SSO 1ue q euonnouny Td 2007 TAU 6 91 0 S pe818 ayi 0 Surpiooog g lt x apqudoosns 8S x gt erpotuoJui x JUB SISOI Y oSewep Jo 9497 2007 TANI IMA ut wraysXg UONEN LAJ wWepurys 0 SUIPIODIY y ynn yayaman oseulep JO OAT Od 05 p OdV 9 18 DOSL dv 106 Od 0 GOV 389 2009 V I6 DEV 00 dod r 89 2009 OdV 28 dv 007 do 109 OAV 00 dogy 8 69 Od 0 dod r 89 Od 0S p qO 7TS9 V 001 q g s cvurp JOON 107 Tdi PIPA IDUBIAOL a OOLST qO L9 9IT qO L9 9IT qO S LI dod SZ SOI dod 0S LOT OadV S7 S9 DIV 0519 V 00 tIl vsLel V ST6I s339 JO N uontsodrao 10 DIOYI ON SOUT IHO PUL SIBAT ND ID ut SNJOIIZI40 SapososN O uustueuo uu JURISTSOI JO UOTFENTRAT TTTAVL x KK NHNHA HH NHN 4 Y oseulep JO PAT av 00 0r V 00 07 av 00 Or av 00 SP av ee 8p V 81 av te eb Od 00 09 D ee 8 Od sr Od 1919 ANELON BATAINS 10 DIFOYI ON sisorqnuy I s Y I Y s Y S Y I I oseurep JO AT av 00 f 35006 av 00 DAV 00 S av 00 Od 0 9 av 00 Od 0 9 VOST av OSE av OSE xP IPO
18. e to the virus Labrin et al 2009 The first one is not cultivated commer cially due to instability grain quality In order to evaluate the genetic constitu tion of commercial varieties and elite lines of the breeding programs of INIA and Danac several experiments were carried out under greenhouse conditions with the aim of char acterizing the different resistance mechanisms to T orizicolus that could be acting Addition ally the tolerance level of cultivars and lines was evaluated MATERIALS AND METHODS Study site Five experiments were estab lished from August 2006 September 2007 in 106 Rev Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 60 1 105 117 March 2012 a greenhouse of Foundation Danac located at 10 21 50 N 685915 W in San Javier Yaracuy State Venezuela The average tem perature and humidity of the greenhouse was 29 5 C and 68 7 HR respectively Rice genotypes Three current commer cial cultivars were evaluated Venezuela 21 Centauro and D Sativa and four promissory advanced lines PNODAO16C PFD 2 1 1 2 gt 293 M 6 2 2 1 FD0241 M 17 6 1 1 1 1 and FD0241 M 18 3 1 2 M 1 the genetic breeding programs of INIA and Fundacion Danac The controls used were Makalioka and Fedearroz 50 as resistant Araure 4 as intermediate and Bluebonnet 50 as suscep tible to sogata Insect populations Insects were collect ed from an insec
19. e us un m todo indirecto para estimar la tolerancia Dentro de los principales par metros determinados se encontraban grado de da o n mero de insectos posados sobre plantas de arroz porcentaje de mortalidad de sogata en estado adulto n mero de huevos y se calcul un ndice de toleran cia con base en la biomasa Los resultados mostraron que los genotipos de arroz evaluados poseen diferentes com binaciones en cuanto a los mecanismos de resistencia as como el grado de su efecto El testigo Bluebonnet 50 se mantuvo consistentemente susceptible en las evaluaciones y el testigo resistente Makalioka fue el nico genotipo que present alta antixenosis y alta antibiosis para sobreviven cia y oviposici n El resto de los genotipos presentaron un menor o mayor grado de resistencia de tipo antixenosis y antibiosis para sobrevivencia y oviposici n Se identific el genotipo FDO241 M 17 6 1 1 1 1 con posible tolerancia al da o directo de sogata Palabras clave arroz sogata virus de la hoja blanca del arroz antixenosis antibiosis tolerancia REFERENCES lvarez R M C E Gamboa M Triana M C Duque amp J Silva 2000 Mecanismo de resistencia a Tagosodes orizicolus Muir Homoptera Delphacidae de tipo antibi tico y no preferencia en algunas l neas de arroz Oryza sativa Investigaci n Agr cola 5 1 Consul tado 28 julio 2008 http www redpavpolar info ve danac viewarticle php id 22 amp layout htm
20. er no choice feeding and susceptibility under free choice It is possible that under free choice D sativa had a heavier dose of infestation than under no choice feeding Antibiosis using no choice test for ovi position These results were similar to those obtained by Pardey et al 1994b who deter mined that Makalioka had higher antibiosis effect for oviposition the same has been report ed for Fedearroz 50 Calvert et al 2004 Morphological differences have been observed in cultivars which have high antibiosis those differences are associated with the presence of pubescent leaves which hamper the oviposi tion of insects or the nutritional quality for the insects Baddi amp Garza 2007 where the absence of some essential amino acid makes the insects weaker to reproduce Heinrichs 1994 Panda amp Khush 1995 Tolerance in greenhouse conditions None of the yield parameters could be evalu ated because plants did not reach the reproduc tive phase due to the direct damage caused by the insects Therefore biomass was reduced during the first growth phases limiting its capacity to recover and produce grain Tolerance to sogata in Araure 4 has been reported previously Vivas 1997 however in this study this cultivar showed a high FPLI and grade six for direct damage these results are consistent with those found under field condition Vivas et al 2002 The cultivar Centauro showed a higher DWI in
21. esistant ones the same result was observed by Pardey et al 1996b Most of the evaluated cultivars showed high to intermediate antibio sis survival actually those genotypes are being used as progenitors in the breeding programs in Venezuela which could increase the possibil ity to select new biotypes of insect Weber amp Parada 1994 Antibiosis should be multigenic characteristic to be stable in rice otherwise the insects would be able to develop new biotypes and races Heinrichs 1994 which could be resistant to antibiosis mechanism from plants Baddii amp Garza 2007 Comparison of direct damage between free choice in row and no choice test The difference observed between free choice and no choice tests is that in the former insects have the option to feed and multiply on the geno types they select whereas in the no choice test the insects are forced to feed on each genotype Pardey et al 1996b The latter is similar to what occurs in commercial planting where one cultivar dominates in which case antixenosis resistance should be short lasting Badii amp Garza 2007 Venezuela 21 and Fedearroz 50 were resistant to insects under free choice feeding however they were susceptible under no choice test Pardey et al 1996b reported very similar results in Fedearroz 50 On the other hand D sativa which is the most plant ed cultivar in Venezuela Fundaci n Danac 2010 exhibited resistance und
22. high insect density conditions showing grade five of direct damage However Centauro showed one of the lowest FPLI values observed among all Rev Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 60 1 105 117 March 2012 113 genotypes According to the method and results of our study we suggest that Centauro is a tolerant cultivar to direct damage caused by T orizicolus Heinrichs et al 1985 Additionally the line FD0241 M 17 6 1 1 1 1 showed indi cations of tolerance although in lower level that Centauro The existence of high antibiosis in Maka lioka interferes with the determination of the tolerance mechanism due to the high mortality of adult insects Thus despite having a FPLI similar to the observed in Centauro the lower level of damage expressed in Makalioka suggests the possibility that the small loss of biomass is due to the early insects death rather than the presence of the tolerance mechanism The graphing analysis proposed by Heinrichs et al 1985 could not be done due to the low number of insects collected at the end of this experiment which allowed the determination of the DWI value Moreover the number of live insects was very low or completely absent at the nymph stage this could be caused by the low number of females at the time of infesta tion since we used insects from fourth and fifth nymphal stages that did not allow differentia tion between males and females
23. l Baddi M H amp V Garza 2007 Resistencia en insectos plantas y microorganismos Universidad Aut noma de Ciudad Ju rez CULCyT A o 4 N 18 Enero febrero Consultado 7 julio 2008 http www2 uacj mx IIT CULCYT EneroFebrero2007 5ARTCULOBADII PDP Calvert L C Pardey M Triana amp R Meneses 2004 Avances en la busqueda de mecanismos de resisten cia a Tagosodes orizicolus Muir Plaga del arroz en Am rica latina Simposio de manejo agroecol gico de plagas en la agricultura tropical Rev Protecci n Veg 19 3 Calvert L Lozano I N Villarreal L Reyes L Romero M Lorieux A Garavito M Duque C Mart nez 2005 Hacia la comprensi n de la base gen tica de la resistencia al complejo virus de la hoja blanca y Tagosodes orizicolus en arroz XLV Reuni n Anual APS San Jos Costa Rica Fundaci n Danac 2010 Fundaci n para la Investigaci n Agr cola Danac Informe Danac 2008 2009 Accio nes para el sector agroalimentario San Javier Danac 63p Consultado 25 mayo 2010 http www danac org ve Fusagri 1995 La sogata Plaga del arroz en el Sistema de Riego Rio Gu rico Fundaci n Servicio Para el Agricultor Noticias Agr colas A o 39 N 14 Marzo abril Heinrichs E A 1994 Biology and management of rice insects Wiley Eastern Limited New Delhi India Heinrichs E A F G Medrano amp H R Rapusas 1985 Genetic evaluation for insect resistance in rice Inter national
24. planting season November April In the central plains area the natural populations of sogata have been charac terized as high since the mid 50 s of past centu ry capturing around 1 700 000 insects per light trap during the dry season Fusagri 1995 Vivas et al 2001 On the other hand the resurgence of populations and consequent losses caused by direct damage have been due to the improper use of pesticides Vivas et al 2007 Vivas et al 2009 It is not known whether there are insect biotypes in Venezuela as it is the case in other countries Hernandez et al 2004 In Venezuela the breeding and release of new rice varieties have been conducted since 1953 by the government agency INIA and by the non governmental sector since the middle of the decade of 1990 Fundaci n Danac is one of these non governmental institutions that have constantly released new technological products contributed to the maintenance of the Venezuelan rice circuit and making more efficient and competitive the rice production Fundaci n Danac 2010 In spite of all the efforts to obtain rice varieties with increased resistance breeding to sogata most of the com mercial varieties in Venezuela present tolerance to direct damage caused by the insect but they do not have a suitable level of resistance to the RHBV Vivas 1997 In the last years only two varieties Fundarroz PN1 and Venezuela 21 have been reported in Venezuela with high level of resistanc
25. rch 2012 117
26. res moleculares microsat lites en arroz Oryza sativa L Gram neae para la resistencia a Tagosodes orizicolus M Homoptera Delphaci dae Tesis Licenciatura Universidad del Valle Cali Colombia Triana M M Cruz R Meneses amp L Calvert 2003 Meto dolog as para la cr a y evaluaci n de Tagosodes ori zicolus Muir Manual T cnico Centro Internacional de Agricultura Tropical Cali Colombia Tambi n disponible en l nea http www ciat cgiar org riceweb index 1 htm Triana M A Pineda C Pardey A Pantoja amp M Duque 1994 Mecanismos de resistencia al da o mec nico por Tagosodes orizicolus Muir en arroz Rev Col Entomol 20 23 26 Vivas L E 1997 Din mica poblacional de la sogata del arroz Tagosodes orizicolus Homoptera Delphaci dae en el Gu rico occidental Tesis de Maestr a Uni versidad Central de Venezuela Maracay Venezuela Vivas L E amp S Clavijo 2000 Fluctuaci n poblacional de Tagosodes orizicolus Muir 1926 Homoptera Delphacidae en el sistema de riego R o Gu rico Calabozo estado Gu rico Venezuela Bol Entomol Venezuela 15 217 227 Vivas L E S Clavijo amp H Gonz lez 2001 Distribuci n temporal y espacial y numero de muestras optimo para la estimaci n de las poblaciones de sogata Tagosodes orizicolus Muir 1926 Homoptera Delphacidae en el cultivo de arroz en Calabozo esta do Gu rico Venezuela Investigaci n Agr cola 6 1 Consultado 1
27. riance ANOVA and mean test using InfoStat program InfoStat 2004 with a significance level of 0 05 When necessary the equivalent non parametric tests were performed RESULTS Antixenosis using free choice test a Circle The ANOVA showed significant differ ences in the number of settled insects between all the genotypes at 24 48 and 72h after infestation p 24h 0 0006 p 48h 0 0003 p 72h 0 0001 The genotypes FD0241 M 17 6 1 1 1 1 Bluebonnet 50 Maka lioka and Araure 4 were more attractive to the insects 24h after infestation Fig 1 However sogata showed less preference to Makalioka at 48 and 72h Bluebonnet 50 and FD0241 M 17 6 1 1 1 1 were the most preferred genotypes since the beginning until the end of the trial The cultivars Araure 4 D Sativa and the line PFD 2111112 gt 293 M 6 2 2 1 showed intermediate antixenosis while FDO241 M 18 3 1 2 M 1 Centauro Makalioka Fedearroz 50 Venezuela 21 and PNOOAO16C showed the highest grade of antixenosis reflected by the low preference of insects b Row The non parametric analy sis detected significant differences between 108 Rev Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 60 1 105 117 March 2012 _25 F X E s EF DE 20 CDEF DEF 3 apcpe BCDEF oe 15 ge ABCDE n ABC ABCD of 10 2 A AB i A l 0 o Se r lt a E 3 E A Y 2 8
28. s Alvarez et al 2002 but under commercial cropping condi tions where the insect density was higher 30 insect double mesh passed this cultivar exhib ited intermediate resistance Fusagri 1995 However it was found that the number of T orizicolus adults in Araure 4 was six fold higher during the dry season in the Sistema Rev Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 60 1 105 117 March 2012 de Riego Rio Gu rico SRRG than the rainy season Vivas amp Clavijo 2000 No report was found comparing simultane ously free choice feeding method circle and row arrangements The daily removal of insects from row method probably decreased the anti xenosis mechanism in some cultivars due to longer exposure of plants to insect effects However row method has been used to evalu ate direct damage in rice breeding programs because it provides a better discrimination among resistant intermediate and susceptible genotypes Pardey et al 1996b Furthermore this method has been useful to study the genet ics of resistance against the direct damage pro duced for sogata Pardey et al 1996a Antibiosis using no choice test for sur vival These results were similar to those found by Triana et al 1994 where Makalioka and Bluebonnet 50 showed the highest and lowest insect mortality respectively Also it was also possible to confirm that T orizicolus lived longer on susceptible cultivars than on r
29. sue with the tip of a syringe needle and using a stereoscope Lieder MZ 720X China 10X to visualize the eggs Alvarez et al 2000 Tolerance in greenhouse conditions Two trials were made for evaluation of tolerance One of them was conducted under a completely randomized design with two replications and the experimental unit EU consisted of two 30 days old plants per pot those plants were covered with micro cages made from 2L clear plastic bottles and infested with a dose of eight insects per plant Other identical trial was performed without insect infestation When the susceptible check Bluebonnet 50 reached nine of the Standard System Evaluation for Rice IRRI 2002 in the infested trial all the plants were removed from the greenhouse infected plants were placed at 20 C for 15 minutes this process allowed to cut down the plants inactivate and remove insect and evaluate the direct damage Subsequently insects and piece of plants with and without infestations were dried at 60 C for 48h Dry weight for insects DWI and dry weight of infested and unin fested plants was registered Subsequently the Functional Plant Loss Index FPLD was calcu lated using the formula proposed by Heinrichs et al 1985 FPLI 1 DWIP DWUP x 1 direct damage 9 x 100 Where DWIP dry weight of infested plants DWUP dry weight of uninfected plants Data from the six experiments were ana lyzed by analysis of va
30. t population at Los Bancos de San Pedro in the Sistema de Riego Rio Gu rico SRRG Gu rico Venezuela locat ed at 8 83 53 N 675340 W where a strong incidence of insects has been reported Fusagri 1995 Laboratory rearing The insects were kept in a cage inside the greenhouse for their reproduction Sequential colonies of T orizi colus free of the RHBV were used Triana et al 2003 Antixenosis using free choice test a Circle adapted from Heinrichs et al 1985 Ten seeds of each genotype and control were planted in pots of 7 50cm high x 9 00cm in diameter A complete randomized block design with four replications was used Plants were thinned to five per pot and fertilized with urea eight days after planting Plants of 25 days were placed into four entomological cages 80cm x 100cm x 65cm in a circular arrangement A dose of five insects per plant of fourth and fifth instar was used and placed in the center of the circle in each cage After 24 48 and 72h of infestation the number of settled insects per plant were counted Finally the direct damage was evaluated 30 days later when Bluebonnet 50 reached grade nine in the range of the Standard System Evaluation for Rice IRRI 2002 The resistance range used was resistant lt 3 intermediated between three and five and susceptible gt 5 b Row adapted from Triana et al 2003 In plastic trays 10 seeds of each material per row at 10cm long and 3
31. xenosis high antibiosis for oviposition medium high antibiosis for survival and tolerance to direct damage Based on antibiosis values it could be a risk to plant this cultivar at a commercial scale in Gu rico State during the dry season when the natural sogata population densities are high Bluebonnet 50 appears to be preferred by the insect for feeding and oviposition it showed low antibiosis for oviposition and high susceptibility to direct damage caused by sogata coinciding with previous reports Pardey et al 1996b Calvert et al 2004 Ara ure 4 behaved similarly to Bluebonnet 50 the difference is that Araure 4 is more tolerant to direct damage making it a good choice for feeding insect mass rearing Under greenhouse conditions the tolerance test allowed the identification of Centauro as possible genotype tolerant to direct damage produced by T orizicolus which should be cor roborated with field trials In summary the tested cultivars expressed a resistance mechanism to sogata differing in the type of mechanism as well as in expres sion level Most commercial rice cultivars tested and elite lines have inadequate antibiotic effects on the insect and they can lead in the future the development of new insect biotypes which can be more aggressive in the produc tion area of the central plains of Venezuela However the line FD0241 M 17 6 1 1 1 1 showed an intermediate behavior for all mech
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