Shading during the grain filling period increases 2-acetyl-1-pyrroline content in fragrant rice

Size: px
Start display at page:

Download "Shading during the grain filling period increases 2-acetyl-1-pyrroline content in fragrant rice"

Transcription

1 Mo et l. Rice (2015) 8:9 DOI /s y RESEARCH Open Access Shding during the grin filling period increses 2-cetyl-1-pyrroline content in frgrnt rice Zhowen Mo 1,2,WuLi 3, Shenggng Pn 1,2, Timothy L Fitzgerld 4, Feng Xio 1, Yongjin Tng 1, Yilei Wng 1, Meiyng Dun 1,2, Hu Tin 1,2 nd Xingru Tng 1,2* Astrct Bckground: Frgrnt rice, including Thi jsmine nd Indin smti vrieties, is highly vlued y consumers glolly. 2-cetyl-1-proline (2-AP) is the mjor compound responsile for the romtic chrcter of frgrnt rice. Previously, environmentl fctors such s wter mngement nd slinity hve een proven to influence 2-AP levels in frgrnt rice; ssessing the effect of dditionl environmentl fctors on 2-AP concentrtion is therefore eminent. The level of solr rdition (solr intensity; SI) to which crop is exposed cn ffect growth, yield nd grin qulity, nd other photosynthetic nd physiologicl chrcteristics. In this study the effect of shding (i.e. the reduction of SI) on yield, qulity, nd 2-AP concentrtion in two elite Chinese frgrnt rice vrieties, Yuxingyouzhn nd Nongxing 18, hs een investigted. Furthermore, ccumultion of the plnt stress response molecules proline nd gmm-minoutyric cid, which hve lso een implicted in pthwys leding to 2-AP production, ws ssessed to study shding effects on these compounds in frgrnt rice, nd to further possily determine fluxes in iochemicl pthwys leding to 2-AP ccumultion. Results: This study hs reveled significnt chnges in the yield nd qulity chrcters under shding tretment. Additionlly, 2-AP nd GABA content in grins ws significntly incresed for ll shding tretments in oth vrieties. In ddition to 2-AP, ten other voltile compounds were studied; results indicted tht shding tretments could hve selective effect on the metolism of these voltile compounds. Conclusions: In this study, we hve demonstrted tht shding during grin filling hs significnt effects on yield nd qulity trits in rice, nd leds to the ccumultion of GABA nd 2-AP. We discuss the implictions of these findings in terms of pthwys leding to 2-AP nd GABA production in frgrnt rice, which hve not een fully elucidted. The shding effect on ten dditionl voltile compounds is lso discussed. Finlly we discuss possile effects of vrition in solr intensity resulting from nthropogenic emissions on frgrnt rice production. Keywords: 2-cetyl-1-pyrroline; Aromtic rice; γ-minoutyric cid; Proline; Shding tretment; Yield Bckground Frgrnt rice vrieties possess chrcteristic rom tht is often descried s nutty or popcorn-like (Brynt nd McClung, 2010). Thi jsmine nd indin smti re the most widely-recognized types of frgrnt rice. Frgrnt rice is highly desired y consumers nd ttrcts premium price in mny mrkets (Skthivel et l., 2009); * Correspondence: Equl contriutors 1 College of Agriculture, South Chin Agriculturl University, Gungzhou, Gungdong, , Chin 2 Scientific Oserving nd Experimentl Sttion of Crop Cultivtion in South Chin, Ministry of Agriculture. P. R. Chin, Gungzhou, Gungdong , Chin Full list of uthor informtion is ville t the end of the rticle furthermore, glol demnd for frgrnt rice is incresing (Hshemi et l., 2013). The voltile compounds in rice re complicted; instrumentl nlyses hve oserved more thn 200 voltile compounds (Tsugit, 1985;Buttery et l., 1988; Chmpgne, 2008). Buttery et l. (1988) demonstrted tht the prole key contriutors to cooked rice rom mong the detected compounds were 2- cetyl-l-pyrroline, (E,E)-2,4-decdienl, nonnl, hexnl, (E)-2-nonenl, octnl, decnl, 4-vinyl-guicol, nd 4-vinylphenol. Jezussek et l. (2002) suggested tht 2-mino cetophenone nd 4,5-epoxy-(E)-2-decenl were importnt previously unknown rice rom compounds. Mrvl et l. (2008) indicted tht 2015 Mo et l.; licensee Springer. This is n Open Access rticle distriuted under the terms of the Cretive Commons Attriution License (http://cretivecommons.org/licenses/y/4.0), which permits unrestricted use, distriution, nd reproduction in ny medium, provided the originl work is properly credited.

2 Mo et l. Rice (2015) 8:9 Pge 2 of 10 orgnic extrcts of cooked romtic rice showed similr rom profile etween the cultivrs, ut showed sustntil differences in the levels of vrious voltile compounds. Moreover, Yng et l. (2008) suggested tht 13 voltile coupounds my contriute to differences in rom. Despite the complexity of voltile compounds in rice (Chmpgne et l., 2005), single voltile compound, 2-cetyl-1-pyrroline (2-AP), is well-proven to e primrily responsile for the romtic chrcter of frgrnt rice (Buttery et l., 1983, 1986; Widjj et l., 1996; Jezussek et l., 2002). In frgrnt rice, 2-AP cn e detected in ll prts of the plnt except the roots (Buttery et l., 1983; Mrvl et l., 2010); 2-AP is lso present in non-frgrnt rice vrieties, ut t much lower concentrtion (Widjj et l., 1996). The inctivtion of BADH2, encoding n minoldehyde dehydrogense, is responsile for the ccumultion of 2-AP in frgrnt rice vrieties (Brdury et l., 2005; Brdury et l., 2008). Proline is precursor of 2-AP in rice (Yoshihshi et l., 2002), nd γ-minoutyrldehyde (GABld) is likely to e the direct precursor to 2-AP production vi BADH2; Brdury et l. (2008) proposed tht functionl BADH2 ctlyses the conversion of GABld to γ-minoutyric cid (GABA), while non-functionl BADH2 in frgrnt rice leds to cyclistion of GABld to Δ 1 -pyrroline, with the ddition of n cetyl group forming 2-AP. The role of proline in plnt stress tolernce is well-estlished (Szdos nd Svouré, 2010); dditionlly, GABA ccumultes in response to iotic nd iotic stresses in plnts (Bouche nd Fromm, 2004) nd is therefore speculted to ply role in stress tolernce (Kinnersley nd Turno, 2000). Thus, it ppers tht there is n overlp in pthwys involved in 2-AP ccumultion nd the response to stress. In further support of n interction etween 2-AP production nd stress responses, environmentl fctors hve een oserved to influence the level of 2-AP in frgrnt rice grins. For exmple, Yoshihshi et l. (2002) reported tht drought stress during grin formtion incresed 2-AP content, nd Gy et l. (2010) nd Poonlphdech et l. (2012) hve lso recently reported n increse in 2-AP ccumultion in response to slt stress. Interestingly, elite frgrnt rice vrieties re reltively susceptile to iotic nd iotic stresses (Niu et l., 2008), nd recently n ssocition of the frgrnce phenotype with slt susceptiility hs lso een reported (Fitzgerld et l., 2010; Wijerthn et l., 2014). Photosyntheticlly Active Rdition (PAR) is the rnge of electromgnetic rdition etween 400 nd 700 nnometers, roughly equivlent to the visile light spectrum, tht plnts (nd most photosynthetic orgnisms) hrness for photosynthesis. The level of PAR to which plnt is exposed is relted to photosynthetic rte (McCree, 1981), nd vrition in PAR cn therefore hve mjor effect on plnt growth nd development. In the field, the intensity of solr rdition (solr intensity; SI) determines the level of PAR to which crop is exposed, nd vrition in solr rdition cn ffect crop yield (Liu nd Tollenr, 2009). Seson, geogrphic loction, nd cloud cover re key fctors influencing SI; dditionlly, SI vries in response to tmospheric pollution nd wter vpour content (Jáuregui nd Luyndo, 1999; Hywood et l., 2011). The level of SI to which plnt is exposed cn lso e strongly ffected y surrounding vegettion; shding stress in plnts growing eneth or mongst cnopy, nd responses to such stress, re well-recognised phenomen (Gommers et l., 2013). To understnd how rice responds to low light environment, reserchers hve used rtificil shding to control light density. Studies hve showed tht shding cn ffect rice morphologicl chrcteristics, physiologicl chrcteristics, yield, nd qulity. For exmple, Tng (1988) found tht shding ffected growth nd yield, decresed dry mtter, nd incresed lef thickness. Ci nd Luo (1999) reported tht shding t different stges decresed rice yield, ccumultion rte of dry mtter nd the uptke of nutrients, ut incresed nutrient content. Ren et l. (2003) indicted tht the distriution nd trnsformtion of plnt dry mtter is ffected y shding, nd some shding levels decresed rice yield. Besides, Deng et l. (2009) found tht the filled grin percentge nd yield were oviously ffected y shding. Moreover, Liu et l. (2012) reported tht the effect of shding on the content of mlondildehyde, soule sugr, soule protein, nd protective enzymes ctivities in leves vrious in genotype. Ding et l. (2004) found tht shding fter heding extended rice growth durtion of the flg lef, decresed decomposing rte of chlorophyll nd MDA content, reduced the photosynthesis of the flg lef, nd tht the degree of the grinfilling ws lower thn tht of under the strong light. Zhng et l. (2007) indicted tht shding ffected rice qulity differently mongst genotypes, however shding incresed protein content nd chlkiness nd decresed mylose content of ll rice cultivrs. Liu et l. (2008) found tht wek light during grin filling ffected rice yield, physiologicl chrcteristics nd qulity, prticulrly the rte of chlky grins. Additionlly, shding hs een demonstrted to effect strch synthse nd relted enzyme ctivities (Wng et l., 2013). Previous studies hve ssessed the effects of n rtificil shding (which results in decrese in SI nd PAR) on growth nd development, yield, nd grin qulity in rice (e.g. Tnk nd Kwno, 1966; Ci nd Luo, 1999; Ren et l., 2003; Zhng et l., 2007). Here we hve studied the impct of shding during the grin filling period on GABA, proline, nd 2-AP ccumultion in frgrnt rice. Furthermore, to provide dditionl insight into the

3 Mo et l. Rice (2015) 8:9 Pge 3 of 10 effect of shding on growth nd yield in frgrnt rice, we hve complemented our nlyses of metolite ccumultion with ssessment of totl nitrogen levels, nd grin yield nd qulity in response to shding tretment. Results Effect of shding tretment on yield, yield relted trits, dry weight, nd hrvest index For Yuxingyouzhn, ll shding tretments resulted in significnt reduction in filled grin percentge, grin weight, grin yield, totl dry weight, nd hrvest index. Additionlly, shding tretment during the whole grin filling period (S1) nd during the ltter stge of grin filling (S3) cused significnt reduction of pnicle numer. For Nongxing 18, shding tretment during the whole grin filling stge S1 nd during erly grin filling (S2) cused significnt reduction of filled grin percentge nd 1000-grin weight; nd S1 significntly decresed grin yield nd totl dry weight (Tle 1). On verge, Yuxingyouzhn hd lower pnicles numer, filled grin percentge, 1000-grin weight, nd totl dry weight, ut hd higher numer of grins per pnicle, grin yield, nd hrvest index thn Nongxing 18 (Tle 1). Effect of shding tretment on grin qulity Significnt effects were identified for some shding tretments on most qulity trits. All shding tretments significntly incresed grin protein content in oth Yuxingyouzhn nd Nongxing 18 (Tle 2). Additionlly, similr effect of shding on chlkiness chrcterstics ws oserved in oth cultivrs; shding during the whole grin filling period nd during lte grin filling resulted in decrese in men rte of chlky grins, nd men degree of chlkiness, however shding during erly grin filling significntly incresed chlkiness chrcteristics (Tle 2). No significnt effect on rown rice rte ws detected in the shding tretments for either cultivr, while for milled rice rte, hed rice rte nd lkli spreding vlue effects vried with timing, during shding s well s cultivr (Tle 2). Effect of shding tretment on 2-cetyl-1-pyrroline, GABA, proline, nd totl nitrogen levels The 2-AP content in grins ws significntly incresed for ll shding tretments in oth vrieties, with increses of %, nd % detected for Yuxingyouzhn nd Nongxing 18, respectively. In oth vrieties, the highest men 2-AP content in grins ws identified for plnts exposed to S2 tretment, with 2-AP contents of μg kg 1 in Yuxingyouzhn nd μg kg 1 in Nongxing 18, compred to respective mens of nd μg kg 1 in the sence of shding (S0) (Figure 1). All shding tretments lso significntly incresed GABA in the grins of Yuxingyouzhn ( % ) nd Nongxing 18 ( %) (Figure 2). In Yuxingyouzhn, significnt increse in proline content ws identified for S1 nd S2, ut not for S3 (Figure 3). For Nongxing 18, proline content ws not significntly ffected y ny shding tretment (Figure 3). No significnt difference in totl nitrogen content in grins etween shding tretments ws oserved (Figure 4). Reltionships mong 2-Acetyl-1-pyrroline, GABA, proline, nd totl nitrogen Significnt correltions were identified etween 2-AP nd GABA in response to shding for Yuxingyouzhn ut Tle 1 Effect of shding tretment on yield, yield relted trits, totl dry weight, nd hrvest index Tretment Pnicles numer (m 2 ) Grins per pnicle Filled grin percentge (%) 1000-grin weight (g) Grin yield (g m 2 ) Totl dry weight(g m 2 ) Yuxingyouzhn Hrvest index (%) S S c c c c S S c c c men * * Nongxing 18 S S c S S men * * Mens in the sme column followed y different lower cse letters for the sme vriety differ significntly t P = 0.05 y LSD tests. Mens of the two vrieties followed y sterisk for the sme detected index difer significnt t P = 0.05 y LSD tests.

4 Mo et l. Rice (2015) 8:9 Pge 4 of 10 Tle 2 Effect of shding tretment on grin qulity Tretment Brown rice rte (%) Milled rice rte (%) Hed rice rte(%) Yuxingyouzhn Protein content(%) Amylose content(%) Alkli grins with chlkiness (%) Chlkiness degree (%) S c c S c c c 7.33 S c S d c 7.70 men 83.67* 72.77* 69.75* 9.85* 24.49* 7.45* 22.08* 10.46* Nongxing 18 S d S c 0.71 c S S c c 0.73 c men Mens in the sme column followed y different lower cse letters for the sme vriety differ significntly t P = 0.05 y LSD tests. Mens of the two vrieties followed y sterisk for the sme grin qulity trit difer significnt t P = 0.05 y LSD tests. none for Nongxing 18. No correltion ws found etween totl nitrogen nd GABA, proline or 2-AP nd etween proline nd totl nitrogen, GABA or 2-AP in either vriety (Tle 3). Effect of shding tretment on the reltive content (%) voltile compounds Eleven voltile compounds were mesured: (E)-2- Hexenl, 1-Hexnol, Heptnl, 2-cetyl-1-pyrroline, Octne, 1-Heptnol,1-Octen-3-ol, Octnl, Benzyl lcohol, Benzenecetldehyde, nd 3,8-Dimethylundecne. For Yuxingyouzhn, shding tretments significntly incresed reltive content in (E)-2-Hexenl. S3 significntly incresed reltive content in 2-AP. Shding tretments (S2 nd S3) significntly decresed reltive content in Octne. S1 tretment significntly incresed the reltive content of Octnl nd Benzyl lcohol. For Nongxing 18, shding tretments (S2 nd S3) significntly incresed the reltive content of (E)-2-Hexenl. Shding tretments significntly incresed the reltive content of 2-AP. S3 significntly decresed the reltive content of Octne, while significntly incresed the reltive content of Octnl. Shding tretment (S1 nd S3) significntly incresed the reltive content of Benzyl lcohol. Shding tretments (S1nd S2) significntly incresed the reltive content of 3,8-Dimethylundecne. As for the comprision etween the two vrieties, Yuxingyouzhn hd significnt higher reltive content of 2-AP thn Nongxing 18. The reltive content of (E)-2-Hexenl, Heptnl, Octne,nd Octnl in Nongxing 18 ws significntly higher thn tht of Yuxingyouzhn (Tle 4). Discussion Shding limits photosyntheticlly ctive rdition (PAR) nd cn e source of plnt stress. Severl studies hve ssessed the effect of shding on growth nd yield in 2-AP content µg kg c Yuxingyouzhn Nongxing 18 Figure 1 Effect of shding tretment on grin 2-AP content in grins. Verticl rs with different lower cse letters ove re significntly different t P = 0.05 y LSD tests. Cpped rs represent SD. d c S0 S1 S2 S3 GABA content µg g Yuxingyouzhn Nongxing 18 Figure 2 Effect of shding tretment on grin GABA content in grins. Verticl rs with different lower cse letters ove re significntly different t P = 0.05 y LSD tests. Cpped rs represent SD. c S0 S1 S2 S3

5 Mo et l. Rice (2015) 8:9 Pge 5 of 10 Proline content µg g c c Yuxingyouzhn Nongxing 18 Figure 3 Effect of shding tretment on proline content in grins. Verticl rs with different lower cse letters ove re significntly different t P = 0.05 y LSD tests. Cpped rs represent SD. cerel crops including rice; shding during flowering nd grin development hs een consistently found to decrese yield (e.g. Beed et l., 2007; Zhng et l., 2006; Liu nd Tollenr, 2009; Deng et l., 2009). Zhng et l. (2007) found tht shding ffects qulity trits in rice, nd generlly, shding resulted in incresed grin protein content nd decresed mylose content, s well s n incresed rte of chlky rice grins nd greter verge degree of chlkinesss. In this study, during the whole (S1), erly (S2), or lte (S3) grin filling period, shding tretment resulted in significntly decresed yield in two elite Chinese frgrnt rice vrieties, with S1 tretment resulting in the gretest yield losses. Yield losses were more sustntil in Yuxingyouzhn thn Nongxing 18 under ll shding tretments; this is consistent with previous studies in which genotype hs een found to influence yield loss due to shding in rice (Zhu et l., 2008; Deng et l., 2009). Shding tretment lso significntly ffected qulity trits. An increse in grin protein content under ll tretments nd consistent effect of tretment on grin chlkiness were oserved for oth vrieties. Significnt shding effects were lso identified for most other qulity trits ssessed, however Totl N content % Yuxingyouzhn Nongxing 18 Figure 4 Effect of shding tretment on totl nitrogen content in grins. Verticl rs with different lower cse letters ove re significntly different t P = 0.05 y LSD tests. Cpped rs represent SD. S0 S1 S2 S3 S0 S1 S2 S3 these vried with timing, durtion of tretment, nd cultivr. Together, these results re lrgely consistent with previous literture nd confirm tht shding during grin development cn hve sustntil effect on grin yield nd qulity in frgrnt rice. Here we hve found tht ll shding tretments significntly incresed the concentrtion of oth 2-AP nd GABA in the grin of Yuxingyouzhn thn Nongxing 18; furthermore, positive correltion etween the ccumultion of these compounds ws oserved in oth vrieties. Although GABA hs een shown to ccumulte in response to rnge of stresses, direct ssessment of the impct of shde on GABA ccumultion ppers limited; our results demonstrte tht the conserved response of GABA to plnt stress extends to shding in rice. Furthermore, our results indicte tht the ssocition etween plnt stress nd ccumultion of 2-AP exists for shding in ddition to drought nd slt. We lso ssessed ccumultion of proline in reponse to shding tretment. This ws significntly incresed in Yuxingyouzhn y S1 nd S2 tretment; in contrst, no significnt effect ws detected for ny shding tretment in Nongxing 18. This suggests tht there my e sustntil vrition of proline ccumultion in response to shding in frgrnt rice vrieties. Our oservtions of the ccumultion of GABA nd 2-AP in response to shding drw ttention to the hypothesized reltionship of the iochemistry of these compounds in frgrnt rice. As outlined ove, in vitro rice BADH2 ctlyses the conversion of GABld to GABA, molecule tht hs een widely implicted in plnt stress responses (Bouche nd Fromm, 2004; Kinnersley nd Turno, 2000) nd it hs een hypothesized tht in frgrnt rice non-functionl BADH2 leds to the ccumultion of GABld, which cyclises nd rects with cetyl group forming 2-AP (Brdury et l., 2008). However, lthough GABA cn e synthesized in plnts vi GABld, the primry pthwy for GABA production is thought to e the GABA shunt, in which glutmte is converted to GABA vi glutmte decroxylse (GAD) (Fit et l., 2008). If 2-AP indeed ccumultes in frgrnt rice vi pthwy tht produces GABA from GABld in the presence of functionl BADH2 (Brdury et l., 2008), it is intriguing tht the ccumultion of oth GABA nd 2-AP increses in response to shde. This suggests tht there my e prllel ctivtion of pthwys leding to GABA in response to shding in frgrnt rice, with 2-AP ccumulting from GABld rther thn GABA due to nonfunctionl BADH2, ut GABA lso ccumulting vi the GABA shunt. However, it is possile tht the ccumultion of oth GABA nd 2-AP occurs vi GABld. If nother enzyme exists in rice tht ctlyses the conversion of GABld to GABA, non-functionl BADH2 my result

6 Mo et l. Rice (2015) 8:9 Pge 6 of 10 Tle 3 Correltion coefficients mong 2-AP, GABA, totl nitrogen, nd proline content in grins Index 2-AP content Grin GABA content Proline content Totl nitrogen content Yuxingyouzhn 2-AP content ** GABA content ** Proline content Totl nitrogen content Nongxing 18 2-AP content GABA content Proline content Totl nitrogen content Significnt correltions t **p < in loss of efficiency rther thn elimintion of GABA ccumultion vi GABld, with excess GABld leding to ccumultion of 2-AP. Brdury et l., (2008) demonstrted tht BADH1, n enzyme with ~ 76% identity to BADH2, ctlyses the formtion of GABA from GABld in vitro, leit t lower efficiency thn BADH2. Furthermore, Singh et l., (2010) identified n ssocition of SNPs predicted to decrese ctivity of BADH1 on GABld with 2-AP-sed rom strength in rice. Thus, this ltter possiility seems plusile. In either cse, modultion of GABA ccumultion in response to stress seems credile explntion for the decrese in stress tolernce reported to e ssocited with frgrnce in rice (Fitzgerld et l., 2010; Wijerthn et l., 2014). For the other voltile compounds in grins, Buttery et l. (1988) demonstrted ten prole key contriutors to cooked rice rom mong the detected compounds. Two of the ten voltile compounds, 2-cetyl-l-pyrroline nd Octnl hve een studied extensively (e.g. Mhtheernont et l., 2001; Jezussek et l., 2002; Wongpornchi et l., 2003; Lohkunjit nd Noomhorm, 2004; Yng et l., 2008; Mrvl et l., 2008; Tnnuwong nd Lertsiri, 2010; Goufo et l., 2011; Goufo et l., 2010; Brynt nd McClung, 2010; Mthure et l., 2014; Mhttntwee nd Rouseff, 2014; Liynrchchi et l., 2014). In our study, 2-AP hd the highest verge reltive content mong the detected compounds in Yuxingyouzhn, while ws the the third most undnt voltile detected in Nongxing 18. Octnl hd the second higher verge reltive content mong the detected compounds in oth vrieties. There ws significnt difference in the verge verge reltive content of the two compounds etween the two vrieties. All shding tretment significntly incresed the reltive content in 2-AP in Nongxing 18, while S3 hd significnt impct on reltive content in 2-AP in Yuxingyouzhn. S1 significntly incresed the reltive content of octnl in oth vrieties. Of the compounds tht were posited to contriute to frgrnce Tle 4 Effect of shding tretment on the reltive content (%) voltile compounds in grins Compounds Yuxingyouzhn Nongxing 18 S0 S1 S2 S3 Men S0 S1 S2 S3 Men (E)-2-Hexenl 0.58c * 1-Hexnol Heptnl * 2-cetyl-1-pyrroline * Octne c 6.01c 6.68* 1-Heptnol Octen-3-ol Octnl * Benzyl lcohol Benzenecetldehyde ,8-Dimethylundecne Mens in the sme row followed y different lower cse letters for the sme vriety differ significntly t P = 0.05 y LSD tests. Mens of the two vrieties followed y sterisk for the sme compound difer significnt t P = 0.05 y LSD tests.

7 Mo et l. Rice (2015) 8:9 Pge 7 of 10 in rice y Yng et l. (2008), in ddition to 2-AP nd octnl, heptnl nd 1-octen-3-ol were lso detected in our studies; nd these two compounds hve lso een reported in other studies (Yng et l., 2008; Tnnuwong nd Lertsiri, 2010; Goufo et l., 2010; Brynt nd McClung, 2010; Mthure et l., 2014). We found tht, heptnl hd the highest verge reltive content in Nongxing 18, nd hd the second high verge reltive content in Yuxingyouzhn. Nongxing 18 hd significnt higher verge reltive content of heptnl thn Yuxingyouzhn. For other voltile compounds, (E)-2- hexenl (Buttery et l.,1988; Yng et l., 2008; Goufo et l., 2010, 2011), 1-hexnol (Mhtheernont et l., 2001; Goufo et l., 2010, 2011; Mthure et l., 2014), octne (Lohkunjit nd Noomhorm, 2004 ), 1-heptnol (Mhtheernont et l., 2001; Yng et l., 2008; Goufo et l., 2010, 2011; Liynrchchi et l., 2014), enzyl lcohol (Wongpornchi et l., 2003; Mthure et l., 2014; Liynrchchi et l., 2014), enzenecetldehyde (Lohkunjit nd Noomhorm, 2004 ), nd 3,8-dimethylundecne (Mhtheernont et l., 2001) were lso detected in our study. Shding incresed the reltive content of (E)-2-hexenl in oth vrieties. There ws no significnt difference etween shding nd non-shding tretments in 1-hexnol, 1-heptnol, nd enzenecetldehyde. Shding (S2 nd S3 for Yuxingyouzhn, S3 for Nongxing 18) decresed the reltive content of octne, nd (S1 for Yuxingyouzhn, S1 nd S3 for Nongxing 18) significntly incresed the reltive content of enzyl lcohol. The significnt difference in 3,8-dimethylundecne ws only found for Nongxing 18 in S1 nd S2 tretments. The results suggest tht shding tretments my hve selective effect on the metolism of the voltile compounds in frgrnt rice. Rice yield, grin qulity nd 2-AP content re key qulities for romtic rice. In this study, we hve ssessed the impct of shding impct on romtic rice yield, grin qulity nd 2-AP content, nd some relted trits. In ddition to providing further insight into shding effects on crop yield nd qulity, nd the interction etween 2-AP ccumultion nd stress in frgrnt rice, the results of this study my hve implictions for rice cultivtion in n environment eing drmticlly influenced y nthropogenic fctors. Air pollution cn hve shding effect, cusing sustntil decreses in surfce level solr intensity (Jáuregui nd Luyndo, 1999; Qin et l., 2006), nd Chin is well-known to experience extreme levels of ir pollution (Chn nd Yo, 2008) including in mjor griculturl regions such s the Yngtze River delt (Sho et l., 2006). There is evidence tht the impct of ir pollution hs decresed nnul solr rdition levels in Gungzhou nd even the whole South Chin region (Liu et l., 2014). Our results suggest tht high pollution levels, prticulrly during grin filling, could influence yield, qulity nd rom in elite Chinese frgrnt rice vrieties. While n incresed content of 2-AP s result of the shding effect of ir pollution my e desirle, yield loss nd loss of grin qulity would clerly e detrimentl. Conclusions Here, we hve demonstrted tht shding during grin filling hs significnt effects on yield nd qulity trits in rice, nd leds to the ccumultion of GABA nd 2-AP. These results further emphsize the importnt reltionship etween stress nd 2-AP production in frgrnt rice. Continuing dvnces in the understnding of this reltionship hve gret potentil for the optimiztion of oth rom production nd stress resistnce in elite rice vrieties. Mterils nd methods Plnt mterils nd tretment conditions Two romtic rice cultivrs, Yuxingyouzhn nd Nongxing 18 were used in this study. These re the min commercil frgrnt rice cultivrs in South Chin. Field experiment ws crried out from July to Novemer, 2013 t South Chin Agriculturl University s Experimentl Frm. This region hs humid sutropicl monsoonl climte. The field consists of sndy, lomy soil nd hs een under pddy cultivtion for mny yers. The min soil properties of the experimentl site were s follows: ph, 4.88; orgnic mtter content, 25.65%; totl N, 1.362%; totl P, 0.958%; totl K, %. The experiment ws rrnged in split-plot design with vrieties s the min plots nd shding tretment s the suplots with three duplictions. After yers of plnting, it is well-known tht the durtion of grin filling in the two vrieties employed for this study is pproximltey 30 dys. Bsed on previous studies, we found tht very strong shding level, corresponding to 90% reduction in full nturl light (s mesured y Luxmeter, model ZDS-10, Chin) resulted in sustntil increse in pest nd diseses dmge. Therefore, we employed shding level equivlent to 67% reduction of full nturl light (s mesured y, Luxmeter, model ZDS-10, Chin) creted y one lyer of lck netting. Four shding tretments were studied: (i) S0: the whole phse during grin filling without shding, tken s the control; (ii) S1: shding during the whole phse of grin filling (30 dys, from Octoer 3rd to Novemer 2nd); (iii) S2: shding during the erly phse of grin filling (15 dys, from Octoer 3rd to Octoer 18th); (iv) S3: shding during the lter phse of the grin filling (15 dys, from Octoer 18th to Novemer 2nd). The whole grin filling stge ws from the end of pollintion to grin dry-down, tht is from R5 to R9 in the report Counce et l. (2000), equivlent to out 30 dys for the two experimentl vrieties. As outlined ove, the whole grin filling stge ws devided

8 Mo et l. Rice (2015) 8:9 Pge 8 of 10 into two stges, one ws the erly phse of grin filling (15 dys), the other ws the lter phse of grin filling (15 dys). The re for ech suplot ws 16 m 2. Seeds were sown on July 15th nd trnsplnting ws performed on August 7th t density of 20 cm 20 cm (out 400 hills per plot) with two seedlings per hill. Plnts were hrvested on Novemer 5th. Fertilizer (1500 kg h 1 ; N, 12.5%; P 2 O 5, 6.0%; K 2 O, 10.0%; orgnic mtter, 15.0%) ws pplied 60% t sl, 40% t tillering stge. Irrigtion, pest nd diseses mngement, nd weed control were the sme in ll tretments following the guidelines recommended y the province. Smpling nd mesurement Determintion of dry weight Plnts from ten rndomly selected hills were tken to the lortory t mturity. Plnts were seprted into leves, (stem + sheth) nd pnicles. Smple were then oven-dried t 80 C to constnt weight for the determintion of dry weight. Determintion of totl nitrogen content Totl nitrogen content in grin ws determined s descried y Lu (1999). Briefly, the dried smples of ground grins (out 0.3 g) were digested using the H 2 SO 4 -HClO 4 method. The digestion ws then used to determine the totl nitrogen content y the Kjeldhl method with 2300 Kjeltec Anlyzer Unit (Foss Tector AB, Sweden). Determintion of yield nd yield relted trits At mturity, grin yield ws mesured from one unit smpling re (1 m 2 ) within ech plot, threshed mnully, then sun dried (djusted to moisture content of ~ 14%). Plnts from ten rndomly selected hills from ech plot were smpled to investigte yield-relted trits. Pnicle numer per m 2 ws mesured y counting the pnicle numers of ech hill within the unit re (1 m 2 )tthree different loctions in ech plot nd the men vlue ws tken s the finl result. Pnicles were threshed mnully, totl numer of grins nd numer of filled grins were counted. Five smples of 1000 grins were tken rndomly from filled seeds, weighted to record 1000-grin weight. Hrvest index ws clculted s the rtio etween grin yield nd dry weight t mturity, expressed s percentge. Determintion of grin qulity Aout 1 kg rice grin from ech tretment ws otined fter storge t room temperture for three months. Brown rice rte ws estimted using rice huller (Jingsu, Chin). Milled rice nd hed rice rtes were estimted using Jingmi testing rice grder (Zhejing, Chin). Percentge of grin with chlkiness nd chlkiness degree were estimted using n SDE-A light ox (Gungzhou, Chin). Amylose content, protein content, nd lkli of grins were determined using n Infrtec 1241 grin nlyzer (FOSS-TECATOR). Determintion of proline content, GABA content nd 2-AP concentrtion For determintion of proline, GABA, nd 2-AP, fresh smple of grins from 1 m 2 re of ech plot were hrvested nd immeditely stored t 20 C. Duplicte mesurements were tken for ech smple nd the men vlue of the three smples per tretment ws tken s the finl result. The proline content ws determined ccording to the method descried y Btes et l. (1973). Briefly, grins (out 0.3 g) ws homogenized in 5 ml of 3% sulfoslicylic cid, then cooled fter heting t oiling wter th for 10 min. Smples filtered nd two ml of the filtrte ws mixed with 3 ml of ninhydrin regent ( 1.25 g ninhydrin in 30 ml glcil cetic cid nd 20 ml 6 M phosphoric cid) nd 2 ml of glcil cetic cid. The rection mixture ws then heted t oiling wter th for 30 min nd plced in n ice th for 20 min efore eing extrcted with 4 ml of toluene. The toluene extrction ws then centrifuged t 4000 rpm for 5 min. The sornce of the red chromophore in the toluene frction ws mesured t 520 nm nd the mount of proline ws determined y comprison with stndrd curve nd expressed s μg g 1. GABA content in grin ws determined s descried y Zho et l. (2009) nd Yo et l. (2008). Briefly, grin (out 0.5 g) ws homogenized in 5 ml of 60% ethnol, treted for 4 hours in oscilltions instrument (HZS-H, Chin) using frequency of 200 oscilltions per minute. The superntnt ws then trnsferred to 5 ml centrifuge tue nd centrifuged t 8000 rpm for 3 min. 1 ml of superntnt ws dded to 10 ml tue, mixed with 0.6 ml 0.2 mol L 1 (ph 9.0) sodium tetrorte, two ml 5% toluene, nd 1 ml 7% sodium hypochlorite, then cooled fter heting t 100 C in wter th for 5 minutes. The sornce of the rection solution ws mesured t 645 nm nd the mount of GABA ws determined y comprison with stndrd curve nd expressed s μg g 1. Grin smples were evluted for 2-AP concentrtion y synchroniztion distilltion nd extrction method (SDE) comined with GCMS-QP 2010 Plus (Shimdzu Corportion, Jpn) s descried y Hung et l. (2012). Briefly, Collidine (2, 4, 6-trimethylpyridine) (Sigm, Switzerlnd), ws used s n internl stndrd. 10 g of finely ground grin ws trnsferred into 500 ml roundottom flsk contining 145 ml purified wter; 5 ml of μg ml 1 internl stndrd ws then dded. A stem

9 Mo et l. Rice (2015) 8:9 Pge 9 of 10 distilltion continuous extrction hed ws ttched to the flsk, nd the flsk ws heted t 150 C y n oil th pot, ZKYY (Gungzhou, Chin). Attched to the other hed of the stem distilltion continuous extrction instrument, diethyl ether (35 ml) ws used s the solvent in 500 ml round-ottom flsk, nd the flsk ws heted t 42 C y wter th pot, HH-2 (Jingsu, Chin). During isoltion, the stem distilltion continuous extrction ws mintined t 10 C y cold wter circultion mchine, YKKY-LX-300 (Beijing, Chin). The isoltion ws performed for 35 min. The ether extrct ws then dried over sodium sulfte, filtered (0.22 μm filter pper, Shimdzu, Jpn), nd then directly used to mesure 2-AP concentrtions with the GCMS-QP 2010 Plus method s descried in Hung et l., (2012). The GCMS- QP 2010 Plus working conditions were s followings: gs chromtogrph equipped with Restek Rxi-5 ms (Shimdzu, Jpn) silic cpillry column (30 m mm μm). The uto injector ws AOC-20i, SPL1. High purity helium gs (99.999%, Gungzhou Gses Co., LTD, Chin) ws the crrier gs t the flow rte of 2.0 ml min 1. The temperture of the GC oven ws 40 C (1 min), incresed t 2 C min 1 to 65 C nd held t 65 C for 1 min, nd then incresed to 220 C t 10 C min 1, nd held t 220 C for 10 min. The ion source temperture ws 200 C. Under these conditions, the retention time of 2-AP ws 7.5 min. 2-AP content ws expressed s μg kg 1.Thereltivecontentofthe rom compounds were identified on the sis of their mssspectrycompringthespectrwiththerecords of the NIST lirry. Sttisticl nlysis Anlysis of vrince nd correltion coefficients were performed using Sttistix 8 (Anlyticl, Tllhssee, Florid, USA). The dt were nlyzed y one-wy nlysis of vrince to ssess differences in yield, yield relted trits, totl dry weight, hrvest index, grin qulity, 2- cetyl-1-pyrroline, GABA, proline nd totl nitrogen etween tretments. Arevitions GABA: γ-minoutyric cid; 2-AP: 2-cetyl-1-pyrroline; GABld: γ-minoutyrldehyde; PAR: Photosyntheticlly Active Rdition; SI: solr intensity. Competing interests The uthors declre no potentil competing interests. Authors contriutions ZM, XT, WL nd SP designed the experiments, ZM,WL nd TLF nlyzed the dt, nd wrote the rticle, FX,YT, YW nd HT performed the trits investigtion, ZM nd MD conducted the GC-MS nlysis. All uthors red nd pproved the finl mnuscript. Acknowledgements This study ws supported y Ntionl Nturl Science Foundtion of Chin ( ), Nturl Science Foundtion of Gungdong Province ( ), Agriculturl Reserch Projects of Gungdong Province (2011AO ), nd Agriculturl Stndrdiztion Project of Gungdong Province (4100 F10003). The uthors would like to thnk Dr. Arydeep Roychoudhury nd Altfhusin B Ndf for their contriutions to the mnuscript. Author detils 1 College of Agriculture, South Chin Agriculturl University, Gungzhou, Gungdong, , Chin. 2 Scientific Oserving nd Experimentl Sttion of Crop Cultivtion in South Chin, Ministry of Agriculture. P. R. Chin, Gungzhou, Gungdong , Chin. 3 Crops Reserch Institute, Gungdong Acdemy of Agriculturl Sciences, Gungzhou, Gungdong , Chin. 4 CSIRO Agriculture Flgship, Queenslnd Bioscience Precinct, 306 Crmody Rd, St Luci, Queenslnd 4067, Austrli. Received: 15 July 2014 Accepted: 8 Jnury 2015 References Btes LS, Wldron RP, Tere ID (1973) Rpid determintion of free proline for wter stress studies. Plnt nd Soil 39: Beed FD, Pveley ND, Sylvester-Brdley R (2007) Predictility of whet growth nd yield in light-limited conditions. J Agric Sci 145(01):63 79 Bouche N, Fromm H (2004) GABA in plnts: just metolite? Trends Plnt Sci 9 (3): Brdury LMT, Fitzgerld TL, Henry RJ, Jin Q, Wters DL (2005) The gene for frgrnce in rice. Plnt Biotechnol J 3(3): Brdury LM, Gillies SA, Brushett DJ, Wters DL, Henry RJ (2008) Inctivtion of n minoldehyde dehydrogense is responsile for frgrnce in rice. Plnt Mol Biol 68(4 5): Brynt RJ, McClung AM (2010) Voltile profiles of romtic nd non-romtic rice cultivrs using SPME/GC MS. Food Chem 124(2): Buttery RG, Ling LC, Mon TR (1986) Quntittive nlysis of 2-cetyl-1-pyrroline in rice. J Agric Food Chem 34(1): Buttery RG, Turnugh JG, Ling LC (1988) Contriution of voltiles to rice rom. J Agric Food Chem 36(5): Buttery RG, Ling LC, Julino BO, Turnugh JG (1983) Cooked rice rom nd 2-cetyl-1-pyrroline. J Agric Food Chem 31(4): Ci KZ, Luo SM (1999) Effect of shding on growth, development nd yield formtion of rice. Chinese Journl of Applied Ecology 10(2): (in Chinese) Chmpgne ET (2008) Rice rom nd flvor: literture review. Cerel Chemistry 85(4): Chmpgne ET, Bett-Grer KL, Thompson J, Mutters R, Grimm CC, McClung AM (2005) Effects of drin nd hrvest dtes on rice sensory nd physicochemicl properties. Cerel Chemistry 82(4): ChnCK,YoX(2008)AirpollutioninmegcitiesinChin.AtmosEnviron42(1):1 42 Counce PA, Keisling TC, Mitchell AJ (2000) A uniform, ojective, nd dptive system for expressing rice development. Crop Sci 40(2): Deng F, Wng L, Yo X, Wng JJ, Ren WJ, Yng WY (2009) Effects of differentgrowing-stge shding on rice grin-filling nd yield. Journl of Sichun Agriculturl University 27(3): (in Chinese) Ding SB, Zhu BY, Wu DY, Zhng L (2004) Effect of temperture nd light on senescence of flg lef nd grin-filling fter rice heding. Journl of South Chin Norml University (Nturl Science Edition) 1: ,128 (in Chinese) Fit A, Fromm H, Wlter D, Glili G, Fernie AR (2008) Highwy or ywy: the metolic role of the GABA shunt in plnts. Trends Plnt Sci 13(1):14 19 Fitzgerld TL, Wters DLE, Brooks LO, Henry RJ (2010) Frgrnce in rice (Oryz stiv) is ssocited with reduced yield under slt tretment. Environmentl nd Experimentl Botny 68(3): Gy F, Mrvl I, Roques S, Gunt Z, Boulnger R, Audeert A, Mestres C (2010) Effect of slinity on yield nd 2-cetyl-1-pyrroline content in the grins of three frgrnt rice cultivrs (Oryz stiv L.) in Cmrgue (Frnce). Field Crop Res 117(1): Gommers CM, Visser EJ, Onge KRS, Voesenek LA, Pierik R (2013) Shde tolernce: when growing tll is not n option. Trends Plnt Sci 18(2):65 71 Goufo P, Dun M, Wongpornchi S, Tng X (2010) Some fctors ffecting the concentrtion of the rom compound 2-cetyl-1-pyrroline in two frgrnt rice cultivrs grown in South Chin. Frontiers of Agriculture in Chin 4(1):1 9 Goufo P, Wongpornchi S, Tng X (2011) Decrese in rice rom fter ppliction of growth regultors. Agronomy for Sustinle Development 31:

10 Mo et l. Rice (2015) 8:9 Pge 10 of 10 Hshemi FSG, Rfii MY, Ismil MR, Mhmud TMM, Rhim HA, Asfliz R, Mlek MA, Ltif MA (2013) Biochemicl, genetic nd moleculr dvnces of frgrnce chrcteristics in rice. Crit Rev Plnt Sci 32(6): Hywood J M, Bellouin N, Jones A, Boucher O, Wild M, Shine K P (2011) The roles of erosol, wter vpor nd cloud in future glol dimming/rightening. Journl of Geophysicl Reserch: Atmospheres ( ), 116(D20203), doi: /2011jd Hung ZL, Tng XR, Wng YL, Chen MJ, Zho ZK, Dun MY, Pn SG (2012) Effects of incresing rom cultivtion on rom nd grin yield of romtic rice nd their mechnism. Scienti Agricultur Sinic 45(6): (in Chinese) Jáuregui E, Luyndo E (1999) Glol rdition ttenution y ir pollution nd its effects on the therml climte in Mexico City. Int J Climtol 19(6): Jezussek M, Julino BO, Schieerle P (2002) Comprison of key rom compounds in cooked rown rice vrieties sed on rom extrct dilution nlyses. J Agric Food Chem 50(5): Kinnersley AM, Turno FJ (2000) Gmm minoutyric cid (GABA) nd plnt responses to stress. Crit Rev Plnt Sci 19(6): Lohkunjit N, Noomhorm A (2004) Supercriticl cron dioxide extrction of 2- cetyl-1-pyrroline nd voltile components from pndn leves. Flvour nd Frgrnce Journl 19(3): Liu AJ, Chen WC, Qin Q (2014) An Anlysis nd estimtion on Solr energy resources in Gungzhou city.gungdong. Meteorology 36(3):59 61 (In Chinese) Liu B, Hn Y, Xie WX, Li JG, Liu J, Go Q (2008) The impct of wek light during grin filling on rice yield, physiology nd qulity. Chinese Rice 5:36 40 (in Chinese) Liu L, Wng L, Deng F, Hung Y, Liu DY, Ren WJ, Yng WY (2012) Osmotic regultion sustnce content nd ctivities of protective enzymes in leves of diffenent hyrid rice comintions s ffected y shding. Chinese Journl of Rice Science 26(5): (in Chinese) Liu W, Tollenr M (2009) Response of yield heterosis to incresing plnt density in mize. Crop Sci 49(5): Liynrchchi GD, Kotterchchi NS, Smrseker R (2014) Voltile profiles of trditionl romtic rice vrieties in Sri Lnk. Journl of the Ntionl Science Foundtion of Sri Lnk 42(1):87 93 Lu RK (1999) Anlys is mesures for gro-chemicl on soil. Chin Agriculturl Science nd Technology Press, Beijing, pp (In Chinese) Mhtheernont S, Kews-rd S, Dumri K (2001) Quntifiction of the rice rom compound, 2-cetyl-1-pyrroline, in uncooked Kho Dwk Mli 105 rown rice. J Agric Food Chem 49(2): Mhttntwee K, Rouseff RL (2014) Comprison of rom ctive nd sulfur voltiles in three frgrnt rice cultivrs using GC Olfctometry nd GC PFPD. Food Chem 154:1 6 Mrvl I, Mestres C, Pernin K, Rieyre F, Boulnger R, Guichrd E, Gunt Z (2008) Odor-ctive compounds in cooked rice cultivrs from Cmrgue (Frnce) nlyzed y GC O nd GC MS. J Agric Food Chem 56(13): Mrvl I, Sen K, Agrei A, Menut C, Morere A, Boulnger R, Gy F, Mestres C, Gunt Z (2010) Quntifiction of 2-cetyl-1-pyrroline in rice y stle isotope dilution ssy through hedspce solid-phse microextrction coupled to gs chromtogrphy tndem mss spectrometry. Anl Chim Act 675(2): Mthure SV, Jwli N, Thengne RJ, Ndf AB (2014) Comprtive quntittive nlysis of hedspce voltiles nd their ssocition with BADH2 mrker in non-smti scented, smti nd non-scented rice ( Oryz stiv L.) cultivrs of Indi. Food Chem 142: McCree KJ (1981) Photosyntheticlly ctive rdition. Physiologicl Plnt Ecology I, Springer Berlin Heidelerg Niu XL, Tng W, Hung WZ, Ren GJ, Wng QL, Luo D, Xio YY, Yng SM, Wng F, Lu BR, Go FY, Lu TG, Liu YS (2008) RNAi-directed downregultion of OsBADH2 results in rom (2-cetyl-1-pyrroline) production in rice (Oryz stiv L.). BMC Plnt Biol 8(1):100. doi: / Poonlphdech J, Mrvl I, Roques S, Audeert A, Boulnger R, Bry X, Gunt Z (2012) Effect of timing nd durtion of slt tretment during growth of frgrnt rice vriety on yield nd 2-Acetyl-1-pyrroline, proline, nd GABA levels. J Agric Food Chem 60(15): Qin Y, Kiser DP, Leung LR, Xu M (2006) More frequent cloud free sky nd less surfce solr rdition in Chin from 1955 to Geophys Res Lett 33(1), L doi: /2005gl Ren WJ, Yng WY, Fn GQ, Zhu XM, M ZH, Xu JW (2003) Effect of low light on dry mtter ccumultion nd yield of rice. Journl of Sichun Agriculturl University 21(4): (in Chinese) Skthivel K, Sundrm RM, Shoh Rni N, Blchndrn SM, Neerj CN (2009) Genetic nd moleculr sis of frgrnce in rice. Biotechnol Adv 27(4): Sho M, Tng XY, Zhng YH, Li WJ (2006) City clusters in Chin: ir nd surfce wter pollution. Front Ecol Environ 4(7): Singh A, Singh PK, Singh R, Pndit A, Mhto AK, Gupt DK, Tygi K, Singh AK, Shrm TR (2010) SNP hplotypes of the BADH1 gene nd their ssocition with rom in rice (Oryz stiv L.). Moleculr reeding 26(2): Szdos L, Svouré A (2010) Proline: multifunctionl mino cid. Trends Plnt Sci 15(2):89 97 Tnk A, Kwno K (1966) Effect of mutul shding on dry-mtter production in the tropicl rice plnt. Plnt nd Soil 24(1): Tnnuwong K, Lertsiri S (2010) Chnges in voltile rom compounds of orgnic frgrnt rice during storge under different conditions. J Sci Food Agric 90(10): Tng YY (1988) Effects of shding on growth, development nd yieldconstituting fctors of rice. Plnt Physiology Communictions 2:50 53 (in Chinese) Tsugit T (1985) Arom of cooked rice. Food Reviews Interntionl 1(3): Wng L, Deng F, Ren WJ, Yng WY (2013) Effects of Shding on strch psting chrcteristics of Indic hyrid rice (Oryz stiv L.). PLoS One 8(7):e doi: /journl.pone Widjj R, Crske JD, Wootton M (1996) Comprtive studies on voltile components of non-frgrnt nd frgrnt rices. J Sci Food Agric 70(2): Wijerthn YMAM, Kotterchchi NS, Gimhni DR, Sirisen DN (2014) Explortion of reltionship etween frgrnt gene nd growth performnces of frgrnt rice (Oryz stiv L.) seedlings under slinity stress. Journl of Experimentl Biology nd Agriculturl Sciences 2(1):7 12 Wongpornchi S, Srisedk T, Choonvisse S (2003) Identifiction nd quntittion of the rice rom compound, 2-cetyl-1-pyrroline, in red flowers (Vllris glr Ktze). J Agric Food Chem 51(2): Yng DS, Shewfelt RL, Lee KS, Kys SJ (2008) Comprison of odor-ctive compounds from six distinctly different rice flvor types. J Agric Food Chem 56(8): Yo S, Yng TW, Zho LJ, Xiong SB (2008) The vrition of γ-minoutyric cid content in germinted rown rice mong different cultivrs. Scienti Agricultur Sinic 41(12): (in Chinese) Yoshihshi T, Kki N, Nguyen TTH, Intomi H (2002) Formtion of flvor compound in romtic rice nd its fluctutions with drought stress. Reserch Highlights JIRCAS 2003(2004):32 33 Zhng JW, Dong ST, Wng KJ, Hu CH, Liu P (2006) Effects of shding on the growth, development nd grin yield of summer mize. Chinese Journl of Applied Ecology 17(4): (in Chinese) Zhng LL, ZhngWZ, Hn YD, Wng ZH, Yn LX, Go SK (2007) Effect of shding on rice qulity of diffenent pnicle types.lioning Agriculturl Science (2): (in Chinese) Zho DW, Pu XY, Zeng YW, Li BX, Du J, Yng SM (2009) Determintion of the γ-minoutyric cid in rley. Journl of Triticee Crops 29(1):69 72 (in Chinese) Zhu P, Yng SM, M J, Li SX, Chen Y (2008) Effect of shding on the photosynthetic chrcteristics nd yield t lter growth stge of hyrid rice comintion. Act Agromomic Sinic 34(11): (in Chinese) Sumit your mnuscript to journl nd enefit from: 7 Convenient online sumission 7 Rigorous peer review 7 Immedite puliction on cceptnce 7 Open ccess: rticles freely ville online 7 High visiility within the field 7 Retining the copyright to your rticle Sumit your next mnuscript t 7 springeropen.com

COVER CROP VARIETY AND SEEDING RATE EFFECTS ON WINTER WEED SEED PRODUCTION

COVER CROP VARIETY AND SEEDING RATE EFFECTS ON WINTER WEED SEED PRODUCTION COVER CROP VARIETY AND SEEDING RATE EFFECTS ON WINTER WEED SEED PRODUCTION Nthn S. Boyd nd Eric B. Brennn, USDA-ARS, Orgnic Reserch Progrm, 1636 E. Alisl Street, Slins, CA 93905 Astrct Weed mngement is

More information

Treatment Spring Late Summer Fall 0.10 5.56 3.85 0.61 6.97 3.01 1.91 3.01 2.13 2.99 5.33 2.50 1.06 3.53 6.10 Mean = 1.33 Mean = 4.88 Mean = 3.

Treatment Spring Late Summer Fall 0.10 5.56 3.85 0.61 6.97 3.01 1.91 3.01 2.13 2.99 5.33 2.50 1.06 3.53 6.10 Mean = 1.33 Mean = 4.88 Mean = 3. The nlysis of vrince (ANOVA) Although the t-test is one of the most commonly used sttisticl hypothesis tests, it hs limittions. The mjor limittion is tht the t-test cn be used to compre the mens of only

More information

Study on enzyme-assisted aqueous extraction of oil from soybean

Study on enzyme-assisted aqueous extraction of oil from soybean 8 Journl Scientific & Industril Reserch J SCI IND RES VOL 69 NOVEMBER 2010 Vol. 69, November 2010, pp. 8-865 Study on enzyme-ssisted queous extrction oil from soyben Jun-Qing Qin*, De-Hui Qin, Xing-Mo

More information

THE PARAMETERS OF TRAPS IN K-FELDSPARS AND THE TL BLEACHING EFFICIENCY

THE PARAMETERS OF TRAPS IN K-FELDSPARS AND THE TL BLEACHING EFFICIENCY GEOCHRONOMETRIA Vol. 2, pp 15-2, 21 Journl on Methods nd Applictions of Asolute Chronology THE PARAMETERS OF TRAPS IN K-FELDSPARS AND THE TL BLEACHING EFFICIENCY ALICJA CHRUŒCIÑSKA 1, HUBERT L. OCZKOWSKI

More information

Small Businesses Decisions to Offer Health Insurance to Employees

Small Businesses Decisions to Offer Health Insurance to Employees Smll Businesses Decisions to Offer Helth Insurnce to Employees Ctherine McLughlin nd Adm Swinurn, June 2014 Employer-sponsored helth insurnce (ESI) is the dominnt source of coverge for nonelderly dults

More information

Effect of viscosity on C sugar in Beet sugar factories

Effect of viscosity on C sugar in Beet sugar factories ANNUAL TRANSACTIONS OF THE NORDIC RHEOLOGY SOCIETY, VOL. 16, 2008 Effect of on C sugr in Beet sugr fctories Mohmmd Hojjtoleslmy 1, Rez Shokrni 2 nd Ahmd Krsi 3 1 Deprtment of food technology, College of

More information

2013 Flax Weed Control Trial

2013 Flax Weed Control Trial 2013 Flx Weed Control Tril Dr. Hether Drby, UVM Extension Agronomist Susn Monhn, Conner Burke, Eric Cummings, nd Hnnh Hrwood UVM Extension Crops nd Soils Technicins 802-524-6501 Visit us on the web: http://www.uvm.edu/extension/cropsoil

More information

** Dpt. Chemical Engineering, Kasetsart University, Bangkok 10900, Thailand

** Dpt. Chemical Engineering, Kasetsart University, Bangkok 10900, Thailand Modelling nd Simultion of hemicl Processes in Multi Pulse TP Experiment P. Phnwdee* S.O. Shekhtmn +. Jrungmnorom** J.T. Gleves ++ * Dpt. hemicl Engineering, Ksetsrt University, Bngkok 10900, Thilnd + Dpt.hemicl

More information

Utilization of Smoking Cessation Benefits in Medicaid Managed Care, 2009-2013

Utilization of Smoking Cessation Benefits in Medicaid Managed Care, 2009-2013 Utiliztion of Smoking Cesstion Benefits in Medicid Mnged Cre, 2009-2013 Office of Qulity nd Ptient Sfety New York Stte Deprtment of Helth Jnury 2015 Introduction According to the New York Stte Tocco Control

More information

The Acoustic Design of Soundproofing Doors and Windows

The Acoustic Design of Soundproofing Doors and Windows 3 The Open Acoustics Journl, 1, 3, 3-37 The Acoustic Design of Soundproofing Doors nd Windows Open Access Nishimur Yuy,1, Nguyen Huy Qung, Nishimur Sohei 1, Nishimur Tsuyoshi 3 nd Yno Tkshi 1 Kummoto Ntionl

More information

Turfgrass and Environmental Research Online

Turfgrass and Environmental Research Online Turfgrss nd Environmentl Reserch Online...Using Science to Benefit Golf USGA s Turfgrss nd Environmentl Reserch Progrm jointly funded coopertive study with GCSAA s Environmentl Institute for Golf to evlute

More information

The Effect of Crumb Rubber Modifier (CRM) on the Performance Properties of Rubberized Binders in HMA pavements

The Effect of Crumb Rubber Modifier (CRM) on the Performance Properties of Rubberized Binders in HMA pavements The Effect of Crum Ruer Modifier (CRM) on the Performnce Properties of Ruerized Binders in HMA pvements Soon-Je Lee* Ph.D. Grdute Student Deprtment of Civil Engineering Clemson University Clemson, SC 29634-0911

More information

Rate and Activation Energy of the Iodination of Acetone

Rate and Activation Energy of the Iodination of Acetone nd Activtion Energ of the Iodintion of Acetone rl N. eer Dte of Eperiment: //00 Florence F. Ls (prtner) Abstrct: The rte, rte lw nd ctivtion energ of the iodintion of cetone re detered b observing the

More information

J4.12 REGIONAL HYDROLOGICAL CYCLE AND WEATHER AND CLIMATE IN THE CONTIGUOUS UNITED STATES

J4.12 REGIONAL HYDROLOGICAL CYCLE AND WEATHER AND CLIMATE IN THE CONTIGUOUS UNITED STATES J4.12 REGIONAL HYDROLOGICAL CYCLE AND WEATHER AND CLIMATE IN THE CONTIGUOUS UNITED STATES 1. INTRODUCTION i Hu 1 nd Song Feng Climte nd Bio-Atmospheric Sciences rogrm School of Nturl Resource Sciences

More information

Enterprise Risk Management Software Buyer s Guide

Enterprise Risk Management Software Buyer s Guide Enterprise Risk Mngement Softwre Buyer s Guide 1. Wht is Enterprise Risk Mngement? 2. Gols of n ERM Progrm 3. Why Implement ERM 4. Steps to Implementing Successful ERM Progrm 5. Key Performnce Indictors

More information

Cooperative Extension Service College of Agricultural, Consumer and Environmental Sciences. Page

Cooperative Extension Service College of Agricultural, Consumer and Environmental Sciences. Page Pest Control in Crops Grown in Northwestern New Mexico, 2014 Annul Dt Report 100-2014 Richrd N. Arnold, Michel K. O Neill, Dniel Smel, Kevin Lomrd, nd Mrgret West 1 Coopertive Extension Service College

More information

DlNBVRGH + Sickness Absence Monitoring Report. Executive of the Council. Purpose of report

DlNBVRGH + Sickness Absence Monitoring Report. Executive of the Council. Purpose of report DlNBVRGH + + THE CITY OF EDINBURGH COUNCIL Sickness Absence Monitoring Report Executive of the Council 8fh My 4 I.I...3 Purpose of report This report quntifies the mount of working time lost s result of

More information

Performance analysis model for big data applications in cloud computing

Performance analysis model for big data applications in cloud computing Butist Villlpndo et l. Journl of Cloud Computing: Advnces, Systems nd Applictions 2014, 3:19 RESEARCH Performnce nlysis model for big dt pplictions in cloud computing Luis Edurdo Butist Villlpndo 1,2,

More information

Helicopter Theme and Variations

Helicopter Theme and Variations Helicopter Theme nd Vritions Or, Some Experimentl Designs Employing Pper Helicopters Some possible explntory vribles re: Who drops the helicopter The length of the rotor bldes The height from which the

More information

EQUATIONS OF LINES AND PLANES

EQUATIONS OF LINES AND PLANES EQUATIONS OF LINES AND PLANES MATH 195, SECTION 59 (VIPUL NAIK) Corresponding mteril in the ook: Section 12.5. Wht students should definitely get: Prmetric eqution of line given in point-direction nd twopoint

More information

Answer, Key Homework 10 David McIntyre 1

Answer, Key Homework 10 David McIntyre 1 Answer, Key Homework 10 Dvid McIntyre 1 This print-out should hve 22 questions, check tht it is complete. Multiple-choice questions my continue on the next column or pge: find ll choices efore mking your

More information

Learner-oriented distance education supporting service system model and applied research

Learner-oriented distance education supporting service system model and applied research SHS Web of Conferences 24, 02001 (2016) DOI: 10.1051/ shsconf/20162402001 C Owned by the uthors, published by EDP Sciences, 2016 Lerner-oriented distnce eduction supporting service system model nd pplied

More information

P.3 Polynomials and Factoring. P.3 an 1. Polynomial STUDY TIP. Example 1 Writing Polynomials in Standard Form. What you should learn

P.3 Polynomials and Factoring. P.3 an 1. Polynomial STUDY TIP. Example 1 Writing Polynomials in Standard Form. What you should learn 33337_0P03.qp 2/27/06 24 9:3 AM Chpter P Pge 24 Prerequisites P.3 Polynomils nd Fctoring Wht you should lern Polynomils An lgeric epression is collection of vriles nd rel numers. The most common type of

More information

2006 IPCC Software for National Greenhouse Gas Inventories: Application and use for India

2006 IPCC Software for National Greenhouse Gas Inventories: Application and use for India 2006 IPCC Softwre for Ntionl Greenhouse Gs Inventories: Appliction nd use for Indi Presenttion for NGGIP Side Event Bonn, June 8, 2013 Prof. Amit Grg (mitgrg@iimhd.ernet.in), INDIA GHG Inventory Softwre

More information

Quality Evaluation of Entrepreneur Education on Graduate Students Based on AHP-fuzzy Comprehensive Evaluation Approach ZhongXiaojun 1, WangYunfeng 2

Quality Evaluation of Entrepreneur Education on Graduate Students Based on AHP-fuzzy Comprehensive Evaluation Approach ZhongXiaojun 1, WangYunfeng 2 Interntionl Journl of Engineering Reserch & Science (IJOER) ISSN [2395-6992] [Vol-2, Issue-1, Jnury- 2016] Qulity Evlution of Entrepreneur Eduction on Grdute Students Bsed on AHP-fuzzy Comprehensive Evlution

More information

CHARACTERIZATION OF HASS AVOCADO PROLEPTIC AND SYLLEPTIC SHOOT PROPORTION IN VALPARAISO REGION, CHILE

CHARACTERIZATION OF HASS AVOCADO PROLEPTIC AND SYLLEPTIC SHOOT PROPORTION IN VALPARAISO REGION, CHILE Proceedings VI World Avocdo Congress (Acts VI Congreso Mundil del Agucte) 2007. Viñ Del Mr, Chile. 12 16 Nov. 2007. ISBN No 978-956-17-0413-8. CHARACTERIZATION OF HASS AVOCADO PROLEPTIC AND SYLLEPTIC SHOOT

More information

Reasoning to Solve Equations and Inequalities

Reasoning to Solve Equations and Inequalities Lesson4 Resoning to Solve Equtions nd Inequlities In erlier work in this unit, you modeled situtions with severl vriles nd equtions. For exmple, suppose you were given usiness plns for concert showing

More information

C-crcs Cognitive - Counselling Research & Conference Services (eissn: 2301-2358)

C-crcs Cognitive - Counselling Research & Conference Services (eissn: 2301-2358) C-crcs Cognitive - Counselling Reserch & Conference Services (eissn: 2301-2358) Volume I Effects of Music Composition Intervention on Elementry School Children b M. Hogenes, B. Vn Oers, R. F. W. Diekstr,

More information

THE EFFECTS OF INCREASED PROTEIN INTAKE ON KIDNEY SIZE AND FUNCTION

THE EFFECTS OF INCREASED PROTEIN INTAKE ON KIDNEY SIZE AND FUNCTION The Journl of Experimentl iology 21, 281 29 (1998) Printed in Gret ritin The Compny of iologists Limited 1998 JE1492 281 THE EFFECTS OF INCRESED PROTEIN INTKE ON KIDNEY SIZE ND FUNCTION KIMERLY. HMMOND

More information

Mateus et al. BMC Biology 2014, 12:97 http://www.biomedcentral.com/1741-7007/12/97

Mateus et al. BMC Biology 2014, 12:97 http://www.biomedcentral.com/1741-7007/12/97 Mteus et l. BMC Biology 2014, 12:97 http://www.iomedcentrl.com/1741-7007/12/97 RESEARCH ARTICLE Open Access Adptive developmentl plsticity: Comprtmentlized responses to environmentl cues nd to corresponding

More information

Basic Analysis of Autarky and Free Trade Models

Basic Analysis of Autarky and Free Trade Models Bsic Anlysis of Autrky nd Free Trde Models AUTARKY Autrky condition in prticulr commodity mrket refers to sitution in which country does not engge in ny trde in tht commodity with other countries. Consequently

More information

Experiment 6: Friction

Experiment 6: Friction Experiment 6: Friction In previous lbs we studied Newton s lws in n idel setting, tht is, one where friction nd ir resistnce were ignored. However, from our everydy experience with motion, we know tht

More information

TITLE THE PRINCIPLES OF COIN-TAP METHOD OF NON-DESTRUCTIVE TESTING

TITLE THE PRINCIPLES OF COIN-TAP METHOD OF NON-DESTRUCTIVE TESTING TITLE THE PRINCIPLES OF COIN-TAP METHOD OF NON-DESTRUCTIVE TESTING Sung Joon Kim*, Dong-Chul Che Kore Aerospce Reserch Institute, 45 Eoeun-Dong, Youseong-Gu, Dejeon, 35-333, Kore Phone : 82-42-86-231 FAX

More information

THERMAL EXPANSION OF TUNGSTEN

THERMAL EXPANSION OF TUNGSTEN . THERMAL EXPANSION OF TUNGSTEN S515 By Peter Hidnert nd W. T. Sweeney ABSTRACT This pper gives the results of n investigtion on the therml expnsion of tungsten (99.98 per cent) over vrious temperture

More information

NQF Level: 2 US No: 7480

NQF Level: 2 US No: 7480 NQF Level: 2 US No: 7480 Assessment Guide Primry Agriculture Rtionl nd irrtionl numers nd numer systems Assessor:.......................................... Workplce / Compny:.................................

More information

Assuming all values are initially zero, what are the values of A and B after executing this Verilog code inside an always block? C=1; A <= C; B = C;

Assuming all values are initially zero, what are the values of A and B after executing this Verilog code inside an always block? C=1; A <= C; B = C; B-26 Appendix B The Bsics of Logic Design Check Yourself ALU n [Arthritic Logic Unit or (rre) Arithmetic Logic Unit] A rndom-numer genertor supplied s stndrd with ll computer systems Stn Kelly-Bootle,

More information

Small Business Networking

Small Business Networking Why Network is n Essentil Productivity Tool for Any Smll Business TechAdvisory.org SME Reports sponsored by Effective technology is essentil for smll businesses looking to increse their productivity. Computer

More information

Economics Letters 65 (1999) 9 15. macroeconomists. a b, Ruth A. Judson, Ann L. Owen. Received 11 December 1998; accepted 12 May 1999

Economics Letters 65 (1999) 9 15. macroeconomists. a b, Ruth A. Judson, Ann L. Owen. Received 11 December 1998; accepted 12 May 1999 Economics Letters 65 (1999) 9 15 Estimting dynmic pnel dt models: guide for q mcroeconomists b, * Ruth A. Judson, Ann L. Owen Federl Reserve Bord of Governors, 0th & C Sts., N.W. Wshington, D.C. 0551,

More information

Small Business Networking

Small Business Networking Why network is n essentil productivity tool for ny smll business Effective technology is essentil for smll businesses looking to increse the productivity of their people nd processes. Introducing technology

More information

Combined Liability Insurance. Information and Communication Technology Proposal form

Combined Liability Insurance. Information and Communication Technology Proposal form Comined Liility Insurnce Informtion nd Communiction Technology Proposl form Comined Liility Insurnce Informtion nd Communiction Technology - Proposl form This proposl form must e completed nd signed y

More information

Homework #6 Chapter 7 Homework Acids and Bases

Homework #6 Chapter 7 Homework Acids and Bases Homework #6 Chpter 7 Homework Acids nd Bses 18. ) H O(l) H 3O (q) OH (q) H 3 O OH Or H O(l) H (q) OH (q) H OH ) HCN(q) H O(l) H 3O (q) CN (q) H 3 O HCN CN Or HCN(q) H (q) CN (q) H CN HCN c) NH 3(q) H O(l)

More information

Small Business Networking

Small Business Networking Why network is n essentil productivity tool for ny smll business Effective technology is essentil for smll businesses looking to increse the productivity of their people nd business. Introducing technology

More information

Small Business Networking

Small Business Networking Why network is n essentil productivity tool for ny smll business Effective technology is essentil for smll businesses looking to increse the productivity of their people nd processes. Introducing technology

More information

ARTICLE IN PRESS. i n t e r n a t i o n a l j o u r n a l o f m e d i c a l i n f o r m a t i c s x x x ( 2 0 1 2 ) xxx xxx

ARTICLE IN PRESS. i n t e r n a t i o n a l j o u r n a l o f m e d i c a l i n f o r m a t i c s x x x ( 2 0 1 2 ) xxx xxx IJB-2938; No. of Pges 12 i n t e r n t i o n l j o u r n l o f m e d i c l i n f o r m t i c s x x x ( 2 0 1 2 ) xxx xxx j ourn l homepge: www.ijmijournl.com Description nd comprison of qulity of electronic

More information

improved (2, 3, 4, 5, 7, 8, 9, 10, 11). Some of the

improved (2, 3, 4, 5, 7, 8, 9, 10, 11). Some of the 8 FLORIDA STATE HORTICULTURAL SOCIETY, 963 EVALUATING SPRAY EQUIPMENT FOR FLORIDA CITRUS R. F. Brooks, W. L. Thompson nd P. T. Jutrs Florid Citrus Experiment Sttion Lke Alfred The first successful, lrge,

More information

2. Transaction Cost Economics

2. Transaction Cost Economics 3 2. Trnsction Cost Economics Trnsctions Trnsctions Cn Cn Be Be Internl Internl or or Externl Externl n n Orgniztion Orgniztion Trnsctions Trnsctions occur occur whenever whenever good good or or service

More information

JaERM Software-as-a-Solution Package

JaERM Software-as-a-Solution Package JERM Softwre-s--Solution Pckge Enterprise Risk Mngement ( ERM ) Public listed compnies nd orgnistions providing finncil services re required by Monetry Authority of Singpore ( MAS ) nd/or Singpore Stock

More information

The LENA TM Language Environment Analysis System:

The LENA TM Language Environment Analysis System: FOUNDATION The LENA TM Lnguge Environment Anlysis System: Audio Specifictions of the DLP-0121 Michel Ford, Chrles T. Ber, Dongxin Xu, Umit Ypnel, Shrmi Gry LENA Foundtion, Boulder, CO LTR-03-2 September

More information

Simulation of operation modes of isochronous cyclotron by a new interative method

Simulation of operation modes of isochronous cyclotron by a new interative method NUKLEONIKA 27;52(1):29 34 ORIGINAL PAPER Simultion of opertion modes of isochronous cyclotron y new intertive method Ryszrd Trszkiewicz, Mrek Tlch, Jcek Sulikowski, Henryk Doruch, Tdeusz Norys, Artur Srok,

More information

The International Association for the Properties of Water and Steam. Release on the Ionization Constant of H 2 O

The International Association for the Properties of Water and Steam. Release on the Ionization Constant of H 2 O The Interntionl Assocition for the Properties of Wter nd Stem Lucerne, Sitzerlnd August 7 Relese on the Ioniztion Constnt of H O 7 The Interntionl Assocition for the Properties of Wter nd Stem Publiction

More information

Small Business Networking

Small Business Networking Why network is n essentil productivity tool for ny smll business Effective technology is essentil for smll businesses looking to increse the productivity of their people nd processes. Introducing technology

More information

Homework #4: Answers. 1. Draw the array of world outputs that free trade allows by making use of each country s transformation schedule.

Homework #4: Answers. 1. Draw the array of world outputs that free trade allows by making use of each country s transformation schedule. Text questions, Chpter 5, problems 1-5: Homework #4: Answers 1. Drw the rry of world outputs tht free trde llows by mking use of ech country s trnsformtion schedule.. Drw it. This digrm is constructed

More information

Module 2. Analysis of Statically Indeterminate Structures by the Matrix Force Method. Version 2 CE IIT, Kharagpur

Module 2. Analysis of Statically Indeterminate Structures by the Matrix Force Method. Version 2 CE IIT, Kharagpur Module Anlysis of Stticlly Indeterminte Structures by the Mtrix Force Method Version CE IIT, Khrgpur esson 9 The Force Method of Anlysis: Bems (Continued) Version CE IIT, Khrgpur Instructionl Objectives

More information

Original Research Accumulation of Pb and Cd and its Effect on Ca Distribution in Maize Seedlings (Zea Mays L.)

Original Research Accumulation of Pb and Cd and its Effect on Ca Distribution in Maize Seedlings (Zea Mays L.) Polish Journl of Environmentl Studies Vol. 14, No. 2 (2005), 203-207 Originl Reserch Accumultion of Pb nd Cd nd its Effect on C Distribution in Mize Seedlings (Ze Mys L.) E. Młkowski 1 *, R. Kurtyk 1,

More information

Unit 29: Inference for Two-Way Tables

Unit 29: Inference for Two-Way Tables Unit 29: Inference for Two-Wy Tbles Prerequisites Unit 13, Two-Wy Tbles is prerequisite for this unit. In ddition, students need some bckground in significnce tests, which ws introduced in Unit 25. Additionl

More information

Evaluation of the urban effect of long-term relative humidity and the separation of temperature and water vapor effects

Evaluation of the urban effect of long-term relative humidity and the separation of temperature and water vapor effects INTERNATIONAL JOURNAL OF CLIMATOLOGY Int. J. Climtol. 27: 1531 1542 (2007) Published online 1 Februry 2007 in Wiley InterScience (www.interscience.wiley.com).1483 Evlution of the urbn effect of long-term

More information

Small Business Networking

Small Business Networking Why network is n essentil productivity tool for ny smll business Effective technology is essentil for smll businesses looking to increse the productivity of their people nd business. Introducing technology

More information

Space Vector Pulse Width Modulation Based Induction Motor with V/F Control

Space Vector Pulse Width Modulation Based Induction Motor with V/F Control Interntionl Journl of Science nd Reserch (IJSR) Spce Vector Pulse Width Modultion Bsed Induction Motor with V/F Control Vikrmrjn Jmbulingm Electricl nd Electronics Engineering, VIT University, Indi Abstrct:

More information

The Velocity Factor of an Insulated Two-Wire Transmission Line

The Velocity Factor of an Insulated Two-Wire Transmission Line The Velocity Fctor of n Insulted Two-Wire Trnsmission Line Problem Kirk T. McDonld Joseph Henry Lbortories, Princeton University, Princeton, NJ 08544 Mrch 7, 008 Estimte the velocity fctor F = v/c nd the

More information

Free Serum Testosterone Level in Male Rats Treated with Tribulus Alatus Extracts

Free Serum Testosterone Level in Male Rats Treated with Tribulus Alatus Extracts Investigtive Urology Triulus Altus Extrcts nd Testosterone Level Interntionl Brz J Urol Vol. 33 (4): 554-559, July - August, 2007 Free Serum Testosterone Level in Mle Rts Treted with Triulus Altus Extrcts

More information

2 DIODE CLIPPING and CLAMPING CIRCUITS

2 DIODE CLIPPING and CLAMPING CIRCUITS 2 DIODE CLIPPING nd CLAMPING CIRCUITS 2.1 Ojectives Understnding the operting principle of diode clipping circuit Understnding the operting principle of clmping circuit Understnding the wveform chnge of

More information

An Undergraduate Curriculum Evaluation with the Analytic Hierarchy Process

An Undergraduate Curriculum Evaluation with the Analytic Hierarchy Process An Undergrdute Curriculum Evlution with the Anlytic Hierrchy Process Les Frir Jessic O. Mtson Jck E. Mtson Deprtment of Industril Engineering P.O. Box 870288 University of Albm Tuscloos, AL. 35487 Abstrct

More information

Utilization of Magnesium Hydroxide Produced by Magnesia Hydration as Fire Retardant for Nylon 6-6,6

Utilization of Magnesium Hydroxide Produced by Magnesia Hydration as Fire Retardant for Nylon 6-6,6 C O M U N I C A Ç Ã O Utiliztion of Mgnesium Hydroxide Produced y Mgnesi Hydrtion s Fire Retrdnt for Nylon 6-6,6 Sôni D.F. Roch Deprtmento de Engenhri Químic, UFMG Virgíni S.T. Ciminelli Deprtmento de

More information

UNIVERSITY OF NOTTINGHAM. Discussion Papers in Economics STRATEGIC SECOND SOURCING IN A VERTICAL STRUCTURE

UNIVERSITY OF NOTTINGHAM. Discussion Papers in Economics STRATEGIC SECOND SOURCING IN A VERTICAL STRUCTURE UNVERSTY OF NOTTNGHAM Discussion Ppers in Economics Discussion Pper No. 04/15 STRATEGC SECOND SOURCNG N A VERTCAL STRUCTURE By Arijit Mukherjee September 004 DP 04/15 SSN 10-438 UNVERSTY OF NOTTNGHAM Discussion

More information

AntiSpyware Enterprise Module 8.5

AntiSpyware Enterprise Module 8.5 AntiSpywre Enterprise Module 8.5 Product Guide Aout the AntiSpywre Enterprise Module The McAfee AntiSpywre Enterprise Module 8.5 is n dd-on to the VirusScn Enterprise 8.5i product tht extends its ility

More information

Folia Forestalia Polonica, series A, 2014, Vol. 56 (2), 79 92 ORIGINAL ARTICLE

Folia Forestalia Polonica, series A, 2014, Vol. 56 (2), 79 92 ORIGINAL ARTICLE Foli Forestli Polonic, series A, 214, Vol. 56 (2), 79 92 ORIGINAL ARTICLE DOI: 1.2478/ffp-214-8 Vriility of selected trits of Ips typogrphus (L.) (Col.: Scolytine) popultions in Beskid ywiecki (Western

More information

Recognition Scheme Forensic Science Content Within Educational Programmes

Recognition Scheme Forensic Science Content Within Educational Programmes Recognition Scheme Forensic Science Content Within Eductionl Progrmmes one Introduction The Chrtered Society of Forensic Sciences (CSoFS) hs been ccrediting the forensic content of full degree courses

More information

Improving Library Users' Perceived Quality, Satisfaction and Loyalty: An Integrated Measurement and Management System

Improving Library Users' Perceived Quality, Satisfaction and Loyalty: An Integrated Measurement and Management System Improving Librry Users' Perceived Qulity, Stisfction nd Loylty: An Integrted Mesurement nd Mngement System by Anne Mrtensen nd Lrs Gr0nholdt This rticle describes the development nd ppliction of structurl

More information

Revisions published in the University of Innsbruck Bulletin of 18 June 2014, Issue 31, No. 509

Revisions published in the University of Innsbruck Bulletin of 18 June 2014, Issue 31, No. 509 Plese note: The following curriculum is for informtion purposes only nd not leglly inding. The leglly inding version is pulished in the pertinent University of Innsruck Bulletins. Originl version pulished

More information

Victims Compensation Claim Status of All Pending Claims and Claims Decided Within the Last Three Years

Victims Compensation Claim Status of All Pending Claims and Claims Decided Within the Last Three Years Claim#:021914-174 Initials: J.T. Last4SSN: 6996 DOB: 5/3/1970 Crime Date: 4/30/2013 Status: Claim is currently under review. Decision expected within 7 days Claim#:041715-334 Initials: M.S. Last4SSN: 2957

More information

Estimating Exchange Rate Exposures:

Estimating Exchange Rate Exposures: Estimting Exchnge Rte Exposures: Issues in Model Structure * Gordon M. Bodnr ** Pul H. Nitze School of Advnced Interntionl Studies, The Johns Hopkins University 1740 Msschusetts Avenue NW Wshington, DC

More information

Examples - What is Ecology? What is Ecology? Systematics Taxonomy. Microbiology. Ecology and Evolution. Physiology. Developmental Biology.

Examples - What is Ecology? What is Ecology? Systematics Taxonomy. Microbiology. Ecology and Evolution. Physiology. Developmental Biology. Wht is Ecology? The study of the interctions between orgnisms nd their environment. Orgnism plnt, niml, microbe Environment biotic Non-living components soil, wter, nutrients Biotic Interctions with other

More information

Operations with Polynomials

Operations with Polynomials 38 Chpter P Prerequisites P.4 Opertions with Polynomils Wht you should lern: Write polynomils in stndrd form nd identify the leding coefficients nd degrees of polynomils Add nd subtrct polynomils Multiply

More information

Performance Monitoring Fundamentals: Demystifying Performance Assessment Techniques

Performance Monitoring Fundamentals: Demystifying Performance Assessment Techniques Performnce Monitoring Fundmentls: Demystifying Performnce Assessment Techniques Roert C. Rice, PhD Rchelle R. Jyringi Dougls J. Cooper, PhD Control Sttion, Inc. Deprtment of Chemicl Engineering Control

More information

Polynomial Functions. Polynomial functions in one variable can be written in expanded form as ( )

Polynomial Functions. Polynomial functions in one variable can be written in expanded form as ( ) Polynomil Functions Polynomil functions in one vrible cn be written in expnded form s n n 1 n 2 2 f x = x + x + x + + x + x+ n n 1 n 2 2 1 0 Exmples of polynomils in expnded form re nd 3 8 7 4 = 5 4 +

More information

Using Bacillus amyloliquefaciens for remediation of aquaculture water

Using Bacillus amyloliquefaciens for remediation of aquaculture water Xie et l. SpringerPlus 13, 2:119 SpringerOpen Journl RESEARCH Open Access Using Bcillus myloliquefciens for remedition of quculture wter Fengxing Xie 2*, Ticheng Zhu 3, Fengfeng Zhng 2, Ke Zhou 2, Yujie

More information

A generic Decision Support System for integrated weed management

A generic Decision Support System for integrated weed management A generic Decision Support System for integrted weed mngement By: Per Rydhl, IPM Consult, Denmrk nd: Nicols Munier-Jolin (INRA), Frnce Robert Msin, University of Pdov, Itly Murizio Sttin (IBAF-CNR), Itly

More information

Version 001 CIRCUITS holland (1290) 1

Version 001 CIRCUITS holland (1290) 1 Version CRCUTS hollnd (9) This print-out should hve questions Multiple-choice questions my continue on the next column or pge find ll choices efore nswering AP M 99 MC points The power dissipted in wire

More information

Key words: extruded, snack, gelatinization, discarded avocados, proximal analysis

Key words: extruded, snack, gelatinization, discarded avocados, proximal analysis Proceedings VI World Avocdo Congress (Acts VI Congreso Mundil del Agucte) 2007. Viñ Del Mr, Chile. 12 16 Nov. 2007. ISBN No 978-956-17-0413-8. USE OF HASS AVOCADO (Perse mericn Mill.) SEED AS A PROCESSED

More information

GFI MilArchiver 6 vs C2C Archive One Policy Mnger GFI Softwre www.gfi.com GFI MilArchiver 6 vs C2C Archive One Policy Mnger GFI MilArchiver 6 C2C Archive One Policy Mnger Who we re Generl fetures Supports

More information

All pay auctions with certain and uncertain prizes a comment

All pay auctions with certain and uncertain prizes a comment CENTER FOR RESEARC IN ECONOMICS AND MANAGEMENT CREAM Publiction No. 1-2015 All py uctions with certin nd uncertin prizes comment Christin Riis All py uctions with certin nd uncertin prizes comment Christin

More information

Growth and Yield of Six Rice Cultivars under Three Water-saving Cultivations

Growth and Yield of Six Rice Cultivars under Three Water-saving Cultivations Plnt Production Science ISSN: 1343-943X (Print) 1349-18 (Online) Journl homepge: http://www.tndfonline.com/loi/tpps2 Growth nd Yield of Six Rice Cultivrs under Three Wter-sving Cultivtions Noki Mtsuo &

More information

Or more simply put, when adding or subtracting quantities, their uncertainties add.

Or more simply put, when adding or subtracting quantities, their uncertainties add. Propgtion of Uncertint through Mthemticl Opertions Since the untit of interest in n eperiment is rrel otined mesuring tht untit directl, we must understnd how error propgtes when mthemticl opertions re

More information

, and the number of electrons is -19. e e 1.60 10 C. The negatively charged electrons move in the direction opposite to the conventional current flow.

, and the number of electrons is -19. e e 1.60 10 C. The negatively charged electrons move in the direction opposite to the conventional current flow. Prolem 1. f current of 80.0 ma exists in metl wire, how mny electrons flow pst given cross section of the wire in 10.0 min? Sketch the directions of the current nd the electrons motion. Solution: The chrge

More information

Econ 4721 Money and Banking Problem Set 2 Answer Key

Econ 4721 Money and Banking Problem Set 2 Answer Key Econ 472 Money nd Bnking Problem Set 2 Answer Key Problem (35 points) Consider n overlpping genertions model in which consumers live for two periods. The number of people born in ech genertion grows in

More information

2015 EDITION. AVMA Report on Veterinary Compensation

2015 EDITION. AVMA Report on Veterinary Compensation 2015 EDITION AVMA Report on Veterinry Compenstion AVMA Report on Veterinry Compenstion 2015 EDITION Copyright 2015 by the All rights reserved. ISBN-13: 978-1-882691-31-9 AVMA Report on Veterinry Compenstion

More information

Health insurance marketplace What to expect in 2014

Health insurance marketplace What to expect in 2014 Helth insurnce mrketplce Wht to expect in 2014 33096VAEENBVA 06/13 The bsics of the mrketplce As prt of the Affordble Cre Act (ACA or helth cre reform lw), strting in 2014 ALL Americns must hve minimum

More information

Assessing authentically in the Graduate Diploma of Education

Assessing authentically in the Graduate Diploma of Education Assessing uthenticlly in the Grdute Diplom of Eduction Dr Mree DinnThompson Dr Ruth Hickey Dr Michelle Lsen WIL Seminr JCU Nov 12 2009 Key ides plnning process tht embeds uthentic ssessment, workintegrted

More information

Distributions. (corresponding to the cumulative distribution function for the discrete case).

Distributions. (corresponding to the cumulative distribution function for the discrete case). Distributions Recll tht n integrble function f : R [,] such tht R f()d = is clled probbility density function (pdf). The distribution function for the pdf is given by F() = (corresponding to the cumultive

More information

WEB DELAY ANALYSIS AND REDUCTION BY USING LOAD BALANCING OF A DNS-BASED WEB SERVER CLUSTER

WEB DELAY ANALYSIS AND REDUCTION BY USING LOAD BALANCING OF A DNS-BASED WEB SERVER CLUSTER Interntionl Journl of Computers nd Applictions, Vol. 9, No., 007 WEB DELAY ANALYSIS AND REDUCTION BY USING LOAD BALANCING OF A DNS-BASED WEB SERVER CLUSTER Y.W. Bi nd Y.C. Wu Abstrct Bsed on our survey

More information

ClearPeaks Customer Care Guide. Business as Usual (BaU) Services Peace of mind for your BI Investment

ClearPeaks Customer Care Guide. Business as Usual (BaU) Services Peace of mind for your BI Investment ClerPeks Customer Cre Guide Business s Usul (BU) Services Pece of mind for your BI Investment ClerPeks Customer Cre Business s Usul Services Tble of Contents 1. Overview...3 Benefits of Choosing ClerPeks

More information

Confirmation by fluorescent tracer of coverage of onion leaves for control of onion thrips using selected nozzles, surfactants and spray volumes

Confirmation by fluorescent tracer of coverage of onion leaves for control of onion thrips using selected nozzles, surfactants and spray volumes RTICLE IN PRESS Crop Protection 26 (27) 1625 1633 www.elsevier.com/locte/cropro Confirmtion y fluorescent trcer of coverge of onion leves for control of onion thrips using selected nozzles, surfctnts nd

More information

Graphs on Logarithmic and Semilogarithmic Paper

Graphs on Logarithmic and Semilogarithmic Paper 0CH_PHClter_TMSETE_ 3//00 :3 PM Pge Grphs on Logrithmic nd Semilogrithmic Pper OBJECTIVES When ou hve completed this chpter, ou should be ble to: Mke grphs on logrithmic nd semilogrithmic pper. Grph empiricl

More information

Vendor Rating for Service Desk Selection

Vendor Rating for Service Desk Selection Vendor Presented By DATE Using the scores of 0, 1, 2, or 3, plese rte the vendor's presenttion on how well they demonstrted the functionl requirements in the res below. Also consider how efficient nd functionl

More information

Matrix Inverse and Condition

Matrix Inverse and Condition Mtrix Inverse nd Condition Berlin Chen Deprtment of Computer Science & Informtion Engineering Ntionl Tiwn Norml University Reference: 1. Applied Numericl Methods with MATLAB for Engineers, Chpter 11 &

More information

Eleni Kalogria Athanasia Varvaresou Spyridon Papageorgiou Evaggelia Protopapa Ioannis Tsaknis Alexios Matikas Irene Panderi

Eleni Kalogria Athanasia Varvaresou Spyridon Papageorgiou Evaggelia Protopapa Ioannis Tsaknis Alexios Matikas Irene Panderi Chromtogrphi (2014) 77:1275 1281 DOI 10.1007/s10337-014-2722-9 ORIGINAL Pre Column Derivtiztion HPLC Procedure for the Quntittion of Aluminium Chlorohydrte in Antiperspirnt Crems Using Quercetin s Chromogenic

More information

Antibody Screening. Antibody Screening in Pre-transfusion Testing and Antenatal Screening

Antibody Screening. Antibody Screening in Pre-transfusion Testing and Antenatal Screening Antiody Screening in Pre-trnsfusion Testing nd Antentl Screening Antiody Screening in Pre-trnsfusion Testing nd Antentl Screening Q. Wht re nturlly occurring or expected ntiodies? Q. Wht re typicl or unexpected

More information