Agronomía Colombiana ISSN: Universidad Nacional de Colombia Colombia

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1 Agronomí Colombin ISSN: Universidd Ncionl de Colombi Colombi Hernández L., Cludi; Hernández G., Mrí Soledd Growth nd development of the cupuçu fruit (Theobrom grndiflorum [Willd. Ex Spreng.] Schum.) in the western colombin Amzon Agronomí Colombin, vol. 3, núm. 1, enero-bril, 1, pp. 95- Universidd Ncionl de Colombi Bogotá, Colombi Avilble in: How to cite Complete issue More informtion bout this rticle Journl's homepge in redlyc.org Scientific Informtion System Network of Scientific Journls from Ltin Americ, the Cribben, Spin nd Portugl Non-profit cdemic project, developed under the open ccess inititive

2 Growth nd development of the cupuçu fruit (Theobrom grndiflorum [Willd. Ex Spreng.] Schum.) in the western colombin Amzon Crecimiento y desrrollo del fruto de copozú (Theobrom grndiflorum [Willd. Ex Spreng.] Schum.) en l Amzoni occidentl colombin Cludi Hernández L. 1, 3 nd Mrí Soledd Hernández G. ABSTRACT Studies of growth nd optiml hrvest time of cupuçu re vitl to ensure fruit qulity nd reduce post-hrvest losses. This studied looked t growth nd fruit development of cupuçu from fruit set to ripening. The mesurements nlyzed included dimeter (longitudinl nd equtoril), fresh nd dry weight, color, ph, titrtble cidity (TA), totl soluble solids (TSS) nd respirtory rte (RR). Three sigmoid sttes were observed during fruit growth: cell division (S1), mximum growth (S), which corresponds to cell expnsion nd growth stbiliztion nd mturtion (S3). The time between fruit set nd physiologicl mturity ws 117 dys. The cupuçu fruit reched physiologicl mturity when it showed chnges in pulp color (H* = 97.1±1.8 ), which coincided with TSS of bout 5.7±.8 Brix, which were good index of mturity long with dys fter fruit set. The respirtion pttern of the cupuçu fruit ws climcteric, with pek of 156.4±4.5 mg CO kg -1 h dys fter fruit set. No ethylene ws detected before hrvest, but ws detected in some fruits posthrvest. Key words: Hue ngle (H*), mturity index, logistic model, respirtory rte. RESUMEN Los estudios de crecimiento y momento óptimo de cosech de copozú son importntes pr grntizr l clidd del fruto y reducir ls pérdids poscosech. Se estudió el crecimiento y desrrollo del fruto de copozú, desde el cujdo hst l mdurez de consumo. Fueron nlizdos diámetro (longitudinl y ecutoril), peso fresco y seco, color, ph, cidez titulble (AT), sólidos solubles totles (SST) e intensidd respirtori (IR). Fueron reconocidos tres estdos tipo sigmoide en el crecimiento del fruto: división celulr (E1), máximo crecimiento (E), el cul corresponde l expnsión celulr, y estbilizción del crecimiento y mdurción (E3). El tiempo trnscurrido entre el cujdo del fruto y l mdurez fisiológic fue 117 dís. El fruto de copozú lcnzó l mdurez fisiológic cundo mostró cmbios en el color de l pulp (H*=97,1±1,8 ), que coincidieron con unos SST lrededor de 5,7±,8 Brix, los cules constituyeron un buen índice de mdurez junto con los dís después del cujdo del fruto. El ptrón respirtorio del fruto de copozú fue de tipo climtérico, con un pico de 156,4±4,5 mg CO kg -1 h dís después del cujdo. No fue detectdo etileno ntes de l cosech, pero si fue detectdo en lgunos frutos en poscosech. Plbrs clve: ángulo Hué (H*), índice de mdurez, modelo logístico, intensidd respirtori. Introduction The cupuçu fruit (Theobrom grndiflorum [Willd. ex Spreng.] Schum.), Fmily Mlvcee (Alverson et l., 1999), hs high economic potentil due to its gribusiness, shown by its high percentge of cidity nd vitmin C in the pulp nd high protein nd ft content in the seed (Crvlho et l., 1999). This species hs thick fleshy fruits (Ibrr-Mnríquez nd Cornejo-Tenorio, ), with hrd epicrp: woody, green skin, covered with dusty, ferruginous lyer (Crvlho et l., 1999). The fruits re usully collected from the ground, s the skin color does not chnge when ripe (Rojs et l., 1998). This prctice cuses dmge due to contmintion nd rodents. For this reson, studies of growth nd optiml hrvest time re importnt to ensure fruit qulity nd reduce post-hrvest losses. Fruits of the genus Theobrom (cupuçu nd mcmbo (T. bicolor)) re clssified s non-climcteric (Crvlho et l., 1999, Hernández et l., 6), however, the posthrvest physiology of cupuçu fruit should be studied further, tking into ccount chemicl, physicl nd sensory developments during the two dys fter hrvest (Crvlho et l., 1999). In the pulp of these fruits t mturtion, the TA Received for publiction: 11 August,. Accepted for publiction: 1 Mrch, Universidd de l Amzoni, Florenci, Cquetá nd Instituto de Cienci y Tecnologí de Alimentos (ICTA), Universidd Ncionl de Colombi. Bogot (Colombi). Instituto Amzónico de Investigciones Científics (Sinchi) nd Instituto de Cienci y Tecnologí de Alimentos (ICTA), Universidd Ncionl de Colombi. Bogot (Colombi). Corresponding uthor. cehernndez@unimzoni.edu.co Agronomí Colombin 3(1), 95-, 1

3 decreses, ph nd TSS increse nd color chnges (from white to cremy yellow in cupuçu, nd light yellow to drk yellow in Mrco) (Hernández et l., 6). In the western colombin Amzon (Cquet), there hve been no studies of growth nd development of cupuçu nd there re no mturity indices. Therefore, the objective of this study ws to nlyze the growth nd development of the fruit; set collection prmeters for cupuçu in the western colombin Amzon, in order to reduce losses due to indequte hrvests nd improve the qulity of the products obtined from the fruit. Mterils nd methods The study ws conducted on the frm Estefni ( N nd W) in Florenci (Cquet), western Colombin Amzon. The environmentl conditions were: ltitude 33 m, men nnul temperture 5.3 C, 86.1% RH, precipittion 3,63.8 mm nd sunshine 1,465.4 h yer -1. The ecotype employed corresponds to ccession A4 from the C.I. Mcgul germplsm bnk. According to Escobr et l. (9), this ecotype hs lrge ovl fruit with shrp pex nd bse; the pulp is yellow. The ripe fruit longitudinl dimeter vries between 16 nd cm (18.7 cm on verge), the equtoril dimeter rnges from.1 to 1. cm (men.58 cm) nd weighs pproximtely 13 g. The 185 recently set fruits were lbeled (4.1±. cm in longitudinl dimeter nd 1.4±.1 cm in equtoril dimeter) from 3 trees on 4.5 yer old plnttion nd were monitored from fruit set until four dys fter nturl bscission. Smples were tken every weeks. The collected fruits were plced in dmp newspper nd trnsported t room temperture in closed plstic bgs to the Nutrition Lbortory, Universidd de l Amzoni (Florenci). The time between collection nd the mesurements ws one hour. Three replictions were observed seprtely for longitudinl nd equtoril dimeter (LD nd ED, respectively) mesured with Vernier cliper,.1 cm precision (model 7-3BPC-6B, Generl Supply Corportion, Jckson, MA). The fresh weight (FW) ws mesured with.1 g precision blnce (model BCC, Precision, Dietikon, Switzerlnd). The dry weight (DW) ws determined by plcing ech of the components of the fruit in n oven t 7 C until constnt weight. The skin color ws mesured t two opposite points long the fruit equtor fter brush removl of the ferruginous lyer, pulp nd seed color were lso mesured. In ll cses, the coordintes L* C* H* were employed with Hunter Lb colorimeter miniscn XE Plus (Illuminnt D65, observer). The respirtory rte (RR) of the fruit ws determined ccording to the sttic method (Kder, ) by confining individul fruits for.5 h t 7 C nd 98,8 kp. The CO nd ethylene production were mesured by gs chromtogrphy using gs chromtogrph (GC) Agilent 489D coupled with n integrtor hp 3395/3396. For mesurements of CO nd C H 4, the GC ws coupled to therml conductivity detector (TCD) nd flme ioniztion detector (FID), respectively. The tempertures of the oven, the injector nd the TCD were 3, 5 nd 3 C, respectively, for the CO determintions. For mesurement of ethylene production, the corresponding tempertures for the oven, the injector nd FID were 3, 5 nd 5 C, respectively. The gs flow rtes (helium, synthetic ir nd hydrogen) were 5-7, 5-6 nd 8-7 m 3 s -1. The column ws clibrted with certified stndrd of % CO nd mol m -3 ethylene (AGA, Bogotá, Colombi). The totl soluble solids (TSS) ws mesured directly from the juice of the pulp of ech fruit with 3 bp portble Atgo refrctometer (Atgo, Jpn). Subsequently, the sme fruit pulp ws homogenized, nd 5 g were mixed with 3 cm 3 of distilled wter, the ph of the resulting mixture ws mesured with Consort C931 electrode electrochemicl nlyzer (Turnhout, Belgium) before being brought to ph of 8.1 with.1 N NOH solution using the titrtble cidity method (TA) (Mercdo- Silv et l., 1998). TA ws reported in percentge by weight of citric cid nd the rtio of TSS nd TA (mturity rtio (MR)) ws tbulted s TSS/citric cid (%). The fruit growth trits were fitted to logistic model: Y = (1) 1 + e b cd Where the coefficient is the mximum reched by the fruit size, b controls the speed of growth, c ffects the slope of the growth curve nd D is the time in dys fter fruit set (DAS). Logistic regression models were estimted using nlyticl softwre Sttistix 9. (Anlyticl Softwre, 8) ccording to Grriz et l. (5) nd Brrer et l. (8). The fitness of the logistic model ws evluted using the vlue of R nd the men squre residul (Grriz et l., 5). Other vribles were subjected to ANOVA with time s fctor of growth, previously checking rndomness, normlity nd homoscedsticity using Sttgrphics Plus (Sttgrphics, ). Mens were compred with the multiple rnges Tukey test, HSD t 95%. 96 Agron. Colomb. 3(1) 1

4 Results Fruit growth Cupuçu s growth conformed to simple sigmoid curve for the dimeters nd fresh weight (Tb. 1, Fig. 1A to 1C). For dry weight, the logistic model chieved good fit (R =.93), but did not present sigmoidl shpe. Mximum growth ws reched bout 83 d fter fruit set. Logistic regression ws good fit for growth trits nd low men squre residul (R.85, P.1). In growth models for fresh nd dry weight three sttes were identified: S1 beginning with fruit set nd where there will be certin number of cell divisions post-set, ccording to sttisticl nlysis S1 lsts up to 33 d. S corresponds to the cell elongtion stge; is from 33 to 83 d. And finl step, S3 corresponding to the stbiliztion step where fruit growth reched its finl size t 96 d (Fig. 1A through 1C). The nturl bscission presented t 139 DAS. The residul degrees of freedom were 58 for the longitudinl nd equtoril dimeters, 118 for fresh weight, 65 for dry weight nd 5 for dry weight of the epicrp, pulp nd seed. The terms were significnt (P.1). The model grows slowly in S1 for FW nd DW (Fig. 1C nd 1D), nd fst for LD nd ED (Fig. 1A nd 1B). The four morphologicl vribles incresed exponentilly during the second stte (S) (Fig. 1A-1D). The fresh weight (FW) ws Tble 1. Logistic models djusted for different vribles vs. growth time (dys fter fruit set) for cupuçu fruits selected ccording to men squre. Dependent vrible Y Logistic model R squre Men residul Longitudinl dimeter (cm) Y = 18.5 / ( 1 + e D ).85.4 Equtoril dimeter (cm) Y =.6 / ( 1 + e D ) Fresh weight (g) Y = / ( 1 + e D ).9 4. Dry weight (g) Y = 5.1 / ( 1 + e D ) Dry weight epicrp (g) Y = 65.4 / ( 1 + e D ).8.5 Dry weight pulp (g) Y = 51.9 / ( 1 + e D ).9. Dry weight seed (g) Y = 5.7 / ( 1 + e D ) less constnt in S3 (Fig. 1C). The longitudinl nd equtoril dimeters nd fresh weight were fitted to simple sigmoid curve with high correltion coefficients (R.85) (Fig. 1A to 1C nd Tb. ) nd sttisticl significnce (P.1). Color The pulp remined white for S1, S nd the beginning of S3 (L* = 75.8±.8, C* = 8.±.4, H* = 98.±1.7 ). During S3, the pulp becme yellow nd intensified the decrese of H * nd the increse of C * until dy 13 (Fig. B). During S3, the pulp presented the lowest H * t dy 143 (9.4±1. ); the highest C * (3.±.7) ws seen t dy 14 (Fig. B). The L* (81.±1.) of the pulp ws highest t dy 14 (Fig. B). Following nturl fruit bscission, L* incresed significntly (P.1) for the 14 d, then declined until dy 143 (Fig. B). Longitudinl dimeter (cm) 3 A Equtoril dimeter (cm) 14 B Fresh weight (g) C Dry weight (g) D Figure 1. Longitudinl (A) nd equtoril (B) dimeters (LD or ED, respectively) nd fresh weight (C) nd dry weight (D) (FW or DW, respectively) during cupuçu fruit growth. The solid line indictes the fitted line of the logistic model. Brs indicte the stndrd error (men ± SE, n = 3 fruits). Hernández L. nd Hernández G.: Growth nd development of the cupuçu fruit (Theobrom grndiflorum [Willd. Ex Spreng.] Schum.) in the western colombin Amzon 97

5 The seed remined similr color to the pulp for S1, S nd the beginning of S3 (L* = 86.8±1., C* = 13.9±1.5, H* = 87.7±1. ). During S3, the seed becme brown with decresing H* until dy 14 (Fig. C). In the seed, H* (47.5±.4 ) nd L* (4.±.1) showed significnt decrese (P.5 nd P.1) t dy 14, with respect to dys 98 nd 117 (Fig. C). The color of the skin beneth the ferruginous lyer becme yellower during S3, nd more intense nd bright with the decrese of H* nd the increse of C* nd L* t dy 14 (Fig. A). For the skin, H*, L* nd C* did not present significnt chnges in S3 (Fig. A). L* ó C* L* ó C* L* ó C* b b b b b b b b 8 b b b 6 b b b > 4 > b Time (dys fter set fruit) FIGURE. Color chnges during cupuçu fruit development S3 in skin (A) pulp (B), nd seeds (C). Brs indicte the stndrd error (men ± SE, n = 3 fruits). Different letters indicte significnt differences ccording to Tukey s test (P.5). b b b b b b b c d cd d d d d d L C H b d H* ( ) H* ( ) H* ( ) A B C Respirtion The RR (mg CO kg -1 h -1 ) ws high (399.37±17.96 mg) in S1 (Fig. 3). Although the beginning of S presented trnsient climcteric, the RR decresed to 97.37±3.3 mg (Fig. 3). Finlly, during S3 pek climcteric ppered (156.4±4.5 mg) t dy 14 (Fig. 3). The increse in the climcteric pek ws significnt (P.5), but did not present detectble ethylene production before nturl fruit bscission. mg CO kg -1 h -1 S S 4 S d b def 15 ef ef f c ef def Time (dys fter set fruit) FIGURE 3. Respirtory rte during cupuçu fruit growth. The brs indicte stndrd error (men ± SE, n = 3 fruits). Different letters indicte significnt differences ccording to Tukey s test (P.5) nd the mesurements of S3 were compred seprtely (d, e, f) to identify the climcteric pek. mg CO kg -1 h b b 16 5 b 14 c 1 15 e d f g c 8 c 6 5 Respirtory rte Ethylene production Time (dys fter set fruit) FIGURE 4. Respirtory rte nd ethylene production of cupuçu fruit posthrvest. Brs indicte the stndrd error (men ± SE, n = 3 fruits). Different letters indicte significnt differences ccording to Tukey s test (P.5). After nturl fruit bscission, the RR incresed from to mg over four dys (Fig. 4), nd presented prllel detectble emission of ethylene (6.3 to L C H 4 kg m -1 h -1 ) (Fig. 4). Other qulity trits During the climcteric pek (14 DAS), TA decresed to.±.%, then incresed to 3.1±.% t the end of S3 (Fig. µl C H 4 kg -1 h Agron. Colomb. 3(1) 1

6 Acidity (% citric cid) ph A.5 Titrtble cidity (%) ph Tiempo (Dís desde el cujdo) TSS (%) MR b B TSS ( Brix) cde cde cde de c e cde b de Mturity rtio Time (dys fter set fruit) Figure 5. Titrtble cidity (TA; % citrus cidt) nd ph (A) nd totl soluble solids (TSS) nd mturity rtio (MR) (B) during S3 cupuçu fruit development. Brs indicte the stndrd error (men ± SE, n = 3 fruits). Different letters indicte significnt differences ccording to Tukey s test (P.5). 5A). The ph showed opposite behvior from TA, with slight increse during the climcteric pek (Fig. 5A), no significnt chnges for either. MR decresed between 98 nd 1 DAS, from 3.5 to.1, then incresed to 3.1 t the end of S3 (Fig. 5B). The TSS showed significnt increse (P.5) 139 DAS from 6.5±1.3 to.5±.6 Brix (Fig. 5B). After nturl bscission, TSS nd MR incresed (.5±.6 to 13.7±.1 Brix nd 3.3 to 4.9, respectively) over d, nd then decresed (Fig. 5B). Discussion Fruit growth The fruit hd single sigmoid growth for 139 d, which coincides with Rojs et l. (1998) for Cquet (14 d) nd Clzvr et l. (1984) for Brzil (-135 d). In contrst, Hernández et l. (6) reported 4 d in Guvire. The difference in development time between northestern Amzon (Guvire) nd western Amzon (Cquet) could be cused by climtic fctors or the existence of genetic diversity (Hernández et l., 7). Models for cupuçu fruit growth, Cquet (Tb. ) differ from those reported by Hernández et l., (6) for Guvire. LD, ED nd FW were fitted to logistic model used for rzá (Hernández et l., 7), Amzonin Ajis (Brrer et l., 8), Abbé Fetel pers (Grriz et l., 5), mong others. According to Brceló et l. (5), the chnge in DW nd FW in S1 is cused by the increse in cell number, likewise the chnge in fruit LD nd is ssocited with the Hernández L. nd Hernández G.: Growth nd development of the cupuçu fruit (Theobrom grndiflorum [Willd. Ex Spreng.] Schum.) in the western colombin Amzon 99

7 cell division process. In the exponentil increse in S, ll morphologicl vribles re ssocited with cell elongtion, ccumultion of reserve photossimiltes nd wter, incresing size of the vcuoles nd the beginning of the ccumultion of orgnic cids, sugrs nd other components. During S3, the fruit dimensions nd FW stbilized, but DW continued to increse (Fig. 1D), ssocited with incresed trnsloction of ssimiltes from the leves. Color None of the color components of the skin chnged significntly during S3. Indicting tht the externl color of the fruit cnnot be used s hrvest index s in bredfruit (Artocrpus ltilis) (Worrell et l., 1998). The pulp color chnge from white to crem yellow during S3 ws lso observed by Hernández et l. (6). Pulp color chnges hve been observed in erly stges of growth nd mturity for other fruits such s mrco (Hernández et l., 6) nd rzá (Glvis nd Hernández, 1993). The C* of the pulp incresed when H* decresed t 13 d. This color behvior hs lso performed in Sweet pepper cv. Domino (Tdesse et l., ). The chnge in color of the pulp my be ssocited with incresed synthesis of crotenoids; cupuçu pulp contins 17.9±4.54 g/ g (Sous et l., 11). In most fruits, conversion of chloroplsts to chromoplsts is ccompnied by synthesis of one or severl kinds of pigments, normlly nthocynins or crotenoids (Hobson, 1999; Kys nd Pull, 4). Importntly, different colors my hve the sme vlue of chrom (C*) nd therefore, this is not n dequte mturity indictor s it is in peppers (López nd Gómez, ). Respirtion The high respirtion rte in erly S1 is due to high degrdtion of substnces through oxide reduction rections, to chieve the necessry energy for the processes of development (growth nd differentition) nd cell mintennce (Brcelo et l., 5; Wills et l., 1998). The respirtory rte t the beginning of S1 is lmost four times lower thn tht reported by Hernández nd Glvis (1994), who used the dynmic method (Kder, ). At the beginning of S, trnsient climcteric ws seen, which hs been observed in Amzonin Ajis (Brrer et l., 8) nd in previous studies on cupuçu (Hernández et l., 6) nd my suggest mechnism tht triggers metbolic processes fter generting chnges ssocited with physiologicl mturity of the fruit (Brrer et l., 8). Climcteric respirtion ws seen fter 14 d (S3) without detectble production of ethylene. This behvior mtches tht observed in guv fruit, cherimoy nd vocdo, where climcteric respirtion significntly precedes incresed ethylene synthesis (Kys nd Pull, 4). The climcteric pek observed is 4.5 times lower thn tht reported by Hernández nd Glvis (1994) for Guvire, which cme 18 d fter fruit set. Tht is, the climcteric pek for cupuçu in Cquetá occurred 56 dys erlier thn in the study on Guvire. This difference is ssocited with shorter growth cycle for cupuçu, Cquet. Cupuçu presents climcteric behvior, result tht contrsts with previous studies tht clssified it s nonclimcteric (Crvlho et l., 1999, Hernández et l., 6). The climcteric pttern mkes it dvisble to hrvest 117 d fter fruit set when the fruit hs reched physiologicl mturity nd the respirtory rte is miniml, just before the climcteric pek. However, this prmeter must be mnged with other, more stble prmeters, tking into ccount the ge of the fruit my vry depending on environmentl fctors nd the cultivr. The respirtory model of cupuçu mtches tht of the pech, which hs similr levels of CO production in the three stges of growth (Seymour et l., 1993). Considering the ctegories proposed by Kder (b), cupuçu cn be clssified s fruit with n extremely high respirtory rte. However, other fruits hve higher respirtory intensities, such s rzá (Eugeni stipitt) (Hernández et l., 7) nd cerol (Mlpighi emrgint) (Crrington nd King, ). The climcteric behvior of cupuçu differs from other fruits of the genus Theobrom such s cco (Kder, b; Kys nd Pull, 4) nd Mrco (Hernández et l., 6), which re clssified s non-climcteric. Ethylene production of cupuçu resembles tht of the pech in the lte S3 stte ( ml C H 4 kg -1 h -1 ) nd kiwi fruit during ripening (6-8 ml C H 4 kg -1 h -1 ) (Seymour et l., 1993). Given the ctegories proposed by Kder (b), cupuçu cn be clssified s fruit with moderte to high ethylene production rte. The increse in the respirtory rte fter bscission my be nturlly ssocited with the processes tht re triggered due to the production of ethylene nd led to senescence. According to the ctegories estblished by Kder (b) nd tking into ccount the dt of extremely high respirtory rte nd high ethylene production, the perishbility Agron. Colomb. 3(1) 1

8 of the cupuçu fruit could be estimted t -4 weeks, but studies should be done in this regrd. Other qulity trits The decrese in TA nd incresed ph during the climcteric pek indicted consumption of orgnic cids s respirtory substrtes. Incresed TA fter this event possibly indictes new synthesis of orgnic cids or n effect on concentrtion by reduced fresh weight (Fig. 1C). The decline in MR between 98 nd 1 d is result of incresed TA nd the stble behvior of TSS before the climcteric pek, becuse TSS incresed significntly only fter this point. This behvior for TA differs from tht previously observed in cupuçu nd mrco (Hernández et l., 6), where TA decresed t the end of S3. The fluctution in TA nd the stble behvior of TSS suggest tht cids re used more thn sugrs for respirtion. The increse in TSS fter the climcteric pek my indicte tht this process triggers conversion mechnism of strch into sugrs. The increse in TSS nd cid t the end of the S3 hs lso been observed in guv (Bulk et l., 1996, Mercdo- Silv et l., 1998) nd feijo (Rodríguez et l., 6). The Cupuçu pulp presented significnt TA nd firly stble but low ph. According to Slisbury nd Ross (6), the low ph my be ssocited with two spects: 1) high cid contents stored in the vcuole nd ) with the growth of the cells, which requires low ph levels. In generl, the incresed ph nd TSS fter the nturl fruit bscission is due to the respirtion process performed to obtin the energy required for metbolic functions; behvior consistent with the results of this study, since s mentioned before, fter bscission of the fruit, respirtory intensity of cupuçu incresed (Fig. C). Conclusions In climtic conditions of the western Colombin Amzon, the cupuçu fruit reches physiologicl mturity 117 dys fter fruit set. When MR is.4. The dimeters nd the fresh weight exhibited single sigmoid growth model. The cupuçu fruit behved s climcteric fruit, with pek of respirtory ctivity t 156.4±4.5 mg CO kg -1 h -1, 14 d fter fruit set. However, no detectble ethylene emission presented pre-hrvest. During S3, the skin color did not chnge significntly, however, the pulp chnged from white to crem yellow, showing tht this chnge cn be used s mturity index. After nturl bscission, the cupuçu fruit decresed ll physicochemicl vribles, RR incresed nd emission of ethylene slowed, but not in ll fruits. The ethylene production levels rnged from 6.3 to ml C H 4 kg -1 h -1 over period of four dys. The prmeters for cupuçu hrvest in the western Colombin Amzon my be the dys fter fruit set (117 d), long with pulp color round H* = 97.1±1.8 nd totl soluble solids vlue of t lest 5.7±.8 Brix. Acknowledgements Thnks to the Instituto Amzónico de Investigciones Científics-Sinchi (Amzonin Scientific Reserch Institute), to Asohofrucol nd the Ministerio de Ambiente, Viviend y Desrrollo Territoril (Ministry of Environment, Housing nd Territoril Development) for the finncil support grnted (1-/6 Project). Literture cited Alverson, W.S., B.A. Whitlock, R. Nyffeler, C. Byer, nd D.A. Bum Phylogeny of the core Mlvles: evidence from sequence dt. Amer. J. Bot. 89, Anlyticl Softwre. 8. STATISTIX for Windows Versión 9. Tllhssee, FL. Brceló, J., R. Nicolás, B. Sbter, nd R. Sánchez. 5. Fisiologí vegetl. Ediciones Pirámide, Mdrid. Brrer, J.A., M.S. Hernández, L.M. Melgrejo, O. Mrtínez, nd J.P. Fernández-Trujillo. 8. Physiologicl behvior nd qulity trits during fruit growth nd ripenning of four Amzonic hot pepper ccessions. J. Sci. Food Agric. 88, Bulk, R.E.E., E.F.E. Bbiker, nd A.H.E. Tiny Chnges in chemicl composition of guv fruits during development nd ripening. Food Chem. 59 (3), Clzvr, B.G., C.H. Müller, nd O. Khwge Fruticultur tropicl: O cupuçuzeiro, cultivo, beneficimento e utilizção do fruto. Embrp; CPATU, Belém, Brzil. Crvlho de, J.E.U., C.H. Müller, R.L. Benchimol, A.K. Kte, nd R.M. Alves Copozú [Theobrom grndiflorum (Willd. Ex Spreng) Shum.]: Cultivo y utilizción. Mnul técnico. Trtdo de Cooperción Amzónic (TCA). Secretrí Protempore Venezuel; FAO; Embrp, Amzoní Orientl, Belem, Brzil. Crrington, C.M.S nd R.A.G. King.. Fruit development nd ripening in Brbdos cherry, Mlpighi emrgint DC. Sci. Hort. 9, 1-7. Escobr, C.J., D. Criollo, nd W. Herrer. 9. Copozú (Theobrom grndiflorum, Willd. Ex Spreng Schum) Vribilidd y mnejo del cultivo en el piedemonte mzónico. Produmedios, Bogot. Glvis, A. nd M.S. Hernández Análisis del crecimiento del fruto y determinción del momento de cosech del rzá (Eugeni stipitt). Revist Colombi Amzónic 6(), Hernández L. nd Hernández G.: Growth nd development of the cupuçu fruit (Theobrom grndiflorum [Willd. Ex Spreng.] Schum.) in the western colombin Amzon 1

9 Grriz, P.I., H.L. Álvrez, nd G.M. Colvit. 5. Growth pttern of Abbé Fetel per fruits. Act Hort. 674, Hernández, M.S. nd A. Glvis Análisis de crecimiento del fruto y determinción del momento de cosech del Copozú. Revist Colombi Amzónic 7(1-), Hernández, M.S., J.A. Brrer, M.P. Crrillo, O. Mrtínez, L.M. Melgrejo, J.A. Glvis, A.E. Css, nd C. Bolños. 6. Crecimiento y desrrollo de los frutos de especies promisoris del género Theobrom, bjo condiciones de l Amzoní norte colombin. pp In: Melgrejo, L.M., M.S. Hernández, J.A. Brrer, nd M.P. Crrillo (eds.). 6. Theobrom. Instituto Amzónico de Investigciones Científics (SINCHI); Universidd Ncionl de Colombi; Ed. Scripto, Bogot. Hernández, M.S., O. Mrtínez, nd J.P. Fernández-Trujillo. 7. Behvior of rzá (Eugeni stipitt Mc Vugh) fruit qulity trits during growth, development nd ripening. Sci. Hort. 111, -7. Hobson, G Mdurción del fruto. pp In: Azcón- Bieto, J. nd M. Tlón (eds.). Fisiologí y bioquímic vegetl. 5 th ed. Intermericn Mc-Grw-Hill, Mdrid. Ibrr-Mnríquez, G. nd G. Cornejo-Tenorio.. Diversidd de frutos de los árboles del bosque tropicl perennifolio de México. Act Bot. Mex. 9, Kder, A.A.. Methods of gs mixing, smpling nd nlysis. pp In: Kder, A.A. (ed.) Posthrvest technology of horticulturl crops. Publ University of Cliforni, Berkeley, CA. Kder, A.A. b. Posthrvest biology nd technology: An overview. pp In: Kder, A.A. (ed.) Posthrvest technology of horticulturl crops. Publ University of Cliforni, Berkeley, CA. Kys, S.J. nd R.E. Pull. 4. Posthrvest Biology. Exon Press, Athens, GA. López, A.F. nd P.A. Gómez.. Developing ripening index for bell peppers bsed on color mesurements. pp In: Artés, F., M.I. Gil, nd M.A. Cones (eds.). Improving posthrvest technologies of fruits, vegetbles nd ornmentls. IIFR Congress. Vol. I. Murci, Spin. Mercdo-Silv, E., P. Benito-Butist, nd M. Grcí-Velsco de A Fruit development, hrvest index nd ripening chnges of guvs produced in centrl Mexico. Posthrv. Biol. Technol. 13, Rodríguez, M., H.E. Arjon, nd H.A. Cmpos. 6. Crcterizción fisicoquímic del crecimiento y desrrollo de los frutos de feijo (Acc sellowin Berg) en los clones 41 (Quimb) y 8-4. Agron. Colomb. 4(1), Rojs, S., J. Zpt, A. Pereir, E. Vrón, C. Crdens, nd F. Cden El cultivo de Copozú (Theobrom grndiflorum) en el piedemonte mzónico colombino. nd ed. Corpoic; Fondo Amzónico, Florenci, Colombi. Slisbury, F.B. nd C.W. Ross. 6. Fisiologí de ls plnts: desrrollo de ls plnts y fisiologí mbientl. Vol. 3. Thompson Editores Spin; Prninfo, Mdrid. Seymour, G.B., J.E. Tylory, nd G.A. Tucker Biochemistry of fruit ripening. Chpmn nd Hll, London. Sttgrphics plus Sttisticl grphics corp. STSC, Rockville, MA. Sous, M.S.B., L.M. Vieir, M.J.M. d Silv, nd A. de Lim. 11. Crcterizção nutricionl e compostos ntioxidntes em resíduos de polps de fruts tropicis. Ciênc. Agrotec. 35(3), Tdesse, T., E. Hewett, M.A. Nichols, nd K.J. Fisher.. Chnges in physicochemicl ttributes of Sweet pepper cv. Domino during fruit growth nd development. Sci. Hort. 93, Wills, R., B. Glsson, D. Grhm, nd D. Joy Posthrvest. An introduction to the physiology nd hndling of fruit, vegetbles nd ornmentls. 4 th ed. CAB, Wllingford, UK. Worrell, D.B., C.M.S. Crrington, nd D.J. Huber Growth, mturtion nd ripening of bredfruit, Artocrpus ltilis (Prk.) Fosb. Sci. Hort. 76, Agron. Colomb. 3(1) 1