Internal breakdown in mango fruit: symptomology and...

Postharvest Biology and Technology 13 (1998) 59–70

Internal breakdown in mango fruit: symptomology andhistology of jelly seed, soft nose and stem-end cavity

Luc Raymond a, Bruce Schaffer a,*, Jeffrey K. Brecht b, Jonathan H. Crane a

a Tropical Research and Education Center, Uni6ersity of Florida, 18905 SW 280 Street, Homestead, FL 33031, USAb Horticultural Sciences Department, P.O. Box 110690, Uni6ersity of Florida, Gaines6ille, FL 32611-0690, USA

Received 14 August 1997; accepted 29 October 1997

Abstract

Fruit of many mango (Mangifera indica L.) cultivars are susceptible to an internal disorder often referred to aseither jelly seed, soft nose, or stem-end cavity (SEC), depending on the symptoms that appear when fruit are cut open.It has not been determined if jelly seed, soft nose, and SEC are different disorders or different symptoms of the samedisorder. Sections of fruit mesocarp from the cultivars Irwin, Tommy Atkins, and Van Dyke were examined by lightmicroscopy to characterize the disorder and discern histological differences or similarities among fruit with each ofthe three types of symptoms. Jelly seed and SEC affected fruit during early fruit ontogeny, whereas soft nosesymptoms were detected only in fully developed fruit. No major microscopic differences were detected among fruitwith jelly seed, soft nose or SEC. In fruit with each type of symptom, disorganization of the cells and rupture of thecell walls were the first microscopic indicators of the disorder, followed by deterioration or dissolution of vascularconnections between the stone and the mesocarp. Stem-end cavity resulted in necrosis of the mesocarp around thecavity. No cavity or tissue necrosis developed around the stone in fruit affected with jelly seed or soft nose. Thepresence of Ca oxalate crystals was observed only in fruit with SEC. Based on temporal and spatial differences insymptom development within the fruit, it appears that soft nose, jelly seed and SEC may be classified as separatedisorders. © 1998 Elsevier Science B.V. All rights reserved.

Keywords: Mango; Internal breakdown; Jelly seed; Soft nose; Stem-end cavity; Histology

1. Introduction

Internal breakdown is used to describe one ormore physiological disorders in mango(Mangifera indica L.) fruit characterized by pre-mature and uneven ripening of the mesocarp. The

* Corresponding author. Tel.: +1 305 2466397 fax: +1 3052467003; e-mail: [email protected]

0925-5214/98/$19.00 © 1998 Elsevier Science B.V. All rights reserved.

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L. Raymond et al. / Posthar6est Biology and Technology 13 (1998) 59–7060

disorder, also known as flesh breakdown or inter-nal flesh breakdown, is referred to as jelly seed,soft nose, or stem-end cavity (SEC). It is notknown whether these terms signify different disor-ders or, as suggested by Winston (1984), differentsymptoms of the same disorder. Most descriptionsin the literature are based on visual observationsof mature and/or ripe fruit. One disadvantagewith such procedures may be that, when the disor-dered fruit can be seen with the naked eye, symp-toms are probably at an advanced stage andconsequently difficult to distinguish. Soft nose hasbeen considered to be similar to tip pulp (Young,1957), or as an advanced stage of flesh breakdown(Malo and Campbell, 1978) or of SEC (Winston,1984). In Indonesia, soft nose and tip pulp areconsidered to be identical to yeasty fruit rot orinsidious fruit rot (Lim and Khoo, 1985). Galan-Sauco et al. (1984) described soft nose as a combi-nation of jelly seed and SEC. The description offlesh breakdown symptoms by Malo and Camp-bell (1978) is very similar to SEC symptoms de-scribed by Mead and Winston (1991).Krishnamurthy (1981) and Subramanayam et al.(1971) used the term internal breakdown to de-scribe the spongy tissue disorder or soft center in‘Alphonso’ mango fruit.

The cause of external breakdown in mangofruit has not been identified. However, Ca defi-ciency has been implicated as the most probablycause (Young, 1957; Young et al., 1962; Burdonet al., 1991), because the incidence of soft nosewas correlated with low Ca concentrations inleaves (Young et al., 1962) and fruit (Burdon etal., 1991).

There is little information in the literatureabout the histology of internal breakdown. It hasbeen reported that cell separation and cell walldegeneration occur in disordered mesocarp offruit affected with soft nose, whereas cell cohesionis maintained in the healthy mesocarp (Burdon etal., 1991). The purpose of this study was toobserve the disorders from their early stages ofdevelopment to fruit maturity, and attempt todiscern histological differences or similaritiesamong jelly seed, soft nose and SEC, and betweendisordered and healthy tissues.

2. Materials and methods

2.1. Collection of samples

Fruit samples were collected from mango(Mangifera indica L.) cvs. Tommy Atkins, Irwin,and Van Dyke trees in an orchard in Homestead,Florida (25.3° N latitude, 80.2° W longitude, 3 mabove sea level) at 2-week intervals in 1995, andonce a week from cv. Tommy Atkins trees in1996, beginning 4 weeks after fruit set (WAFS)and continuing until the fruit were ripe. Fruitwere considered ripe or mature when theyattained maximum weight and the mesocarpwas soft enough for human consumption (14WAFS).

To assess the type of disorder, one longitudinalcut was made at the proximal end of the fruit,immediately under the epidermis, to expose thetissues in the peduncular extension and to assessthe presence of SEC. The term peduncular exten-sion is used as a synonym of proximal end to referto the vascular connections located between thepeduncle and the stone of the mango fruit, whereSEC symptoms generally develop. Two lateralcuts were made on each flat side of the stone toexpose the interior of the fruit so that jelly seed orsoft nose could be detected. The symptoms werevisually rated as early, intermediate, or advanced.For SEC, rating of the symptoms was based onthe presence and/or the size of the cavity at theproximal end of the fruit. Early symptoms of SECwere those in which no visible cavity was formedat the peduncular extension of the fruit, but wherethe vascular tissues in that area had partly orcompletely turned a brownish color. Symptoms ofSEC were considered as intermediate when a 0.1–0.5 cm diameter cavity was visible at the proximalend of the fruit, accompanied by a necrotic areaaround that cavity. Symptoms were ranked asadvanced when the cavity was larger than 0.5 cmdiameter and was accompanied by deteriorationof the fruit mesocarp around the stone.

For jelly seed and soft nose, early symptomswere characterized by a pronounced yellow col-oration of the mesocarp around the stone (jellyseed) or at the distal end or at the sinus of thefruit (soft nose), indicating that the senescence

L. Raymond et al. / Posthar6est Biology and Technology 13 (1998) 59–70 61

process had been initiated. Intermediate symp-toms were those in which the disordered mesocarphad an orange color, whereas the surroundingtissue (exterior of the mesocarp) was still white orpale yellow, indicating that the exterior of themesocarp was not ripe. Symptoms of jelly seedand soft nose were considered as advanced if themesocarp showed advanced deterioration and/ordiscoloration. All fruit showing a combination ofSEC and jelly seed symptoms were scored as SEC,and those with a combination of jelly seed andsoft nose were scored as jelly seed.

Tissues sections (0.5×0.5×0.75 cm) of thepeduncular extension, the interior of the meso-carp, or at the distal end of the fruit were excisedfrom both healthy and disordered fruit. In fruitwith internal breakdown, samples were taken onlyfrom the disordered areas. Excised samples wereimmediately placed in fixative (McDowell andTrump, 1976).

2.2. Infiltration and embedding

The excised samples from the peduncular exten-sion of the fruit and the mesocarp remained in thefixative for 12–16 weeks. The samples werewashed in 25% EtOH and then placed in 30%EtOH for 2 h and then 50% EtOH for 2 h. Afterair was removed from the samples in a vacuumoven, they were transferred to 70% tertiary butylalcohol (TBA) for 10 h, followed by 85 and 95%TBA each for 2 h. The tissue samples were thensuccessively transferred three times to 100% TBAfor 2, 2, and 1 h, respectively. Finally, the sampleswere placed in another 100% TBA bath for a 10-hperiod. The samples were removed from the TBAand placed into a 1:1 mixture of TBA plus min-eral oil for 2 h. Next, the samples were embeddedin paraffin as described by Johansen (1940).

2.3. Microtomy

Embedded tissues were affixed on 2×2×2 cmwooden blocks and sectioned with a rotary micro-tome to 8–12 mm thickness, depending on thecondition of the tissue. For fruit with SEC, ex-cised samples from the peduncular extension weresectioned both transversely and longitudinally.

Tissue sections were fastened to glass slides withadhesive prepared by dissolving 1 g of high gradeKnox gelatin in 100 ml of deionized water at35°C. After the gelatin was completely dissolved,5 g of phenol and 15 ml of glycerin were addedand the preparation was combined with 3% for-malin (T. Lucansky, University of Florida, per-sonal communication).

2.4. Staining

The paraffin was removed by placing the slidesin glass staining dishes containing 100% xylene,twice for 5 min each time. The slides were thentransferred to 1:1 xylene and EtOH for 5 min,successively transferred to 100, 95, 70, 50, and30% EtOH for 5 min each, and stained with a 1%Safranin-O (Fischer Scientific, Pittsburgh, PA) so-lution for 8 h. After the Safranin was removedwith distilled water and successive EtOH dips, thetissue sections were stained with Fast Green FCF.The Fast Green solution was prepared by dissolv-ing 1 g of Fast Green FCF (Fischer Scientific,Pittsburgh, PA, USA) in a mixture of 125 mlclove oil plus 125 ml 95% EtOH (B. Dehgan,University of Florida, personal communication).The slides were then successively transferred to 95and 100% EtOH, followed by a 1:1 xylene and100% EtOH treatment for 3 min. Finally, theslides were placed in three consecutive baths of100% xylene for 3 min each time and mounted inPermount (Fischer Scientific, Pittsburgh, PA,USA). A WILD MPS52 camera (Leitz, Heerburg,Switzerland) connected to a Laborlux S micro-scope (Leitz, Heerburg, Switzerland) at 10–20×magnification was used to photograph the pre-pared sections.

3. Results

Soft nose can be described as a partial ripeningof the mesocarp at the distal end of the fruit,which in its early stage results in a defined yellowarea between the apex of the stone and the exo-carp (Fig. 1a). Jelly seed affects the interior of themesocarp and is first characterized by a morepronounced yellow color of the affected area com-


Fig. 1. Visual symptoms of a) early stage of soft nose, b) advanced stage of jelly seed and c) advance stage of stem-end cavity (SEC)in mango fruit.

pared to the rest of the mesocarp, which remainswhitish or pale green in young fruit. As the symp-toms develop, the yellow color intensifies andreaches a larger portion of the mesocarp. Theaffected area eventually becomes brown and soft-ens to the point of having the consistency of jelly(Fig. 1b). Stem-end cavity is characterized by theformation of a cavity in the proximal area of thefruit resulting from the deterioration of the vascu-lar tissues between the proximal end of the stoneand the fruit peduncle. The affected tissues turn abrownish color at the earliest stage; then a smallcavity forms as necrosis develops around the cav-ity. The interior of the mesocarp turns yellow ororange, whereas the exterior of the mesocarp re-mains whitish or pale yellow. The cavity laterenlarges as the fruit reaches the advanced stagesof the disorder (Fig. 1c).

The time at which the first symptoms of inter-nal breakdown appeared varied with the cultivars

and with the type of symptoms (Table 1). Stemend cavity and jelly seed were first observed 8WAFS in cvs. Tommy Atkins and Van Dyke,when the average fruit weight was approximately17 and 10% of the final weight of each cultivar,

Table 1Time at which jelly seed, stem-end cavity (SEC) and soft nosedisorders were first detected in ‘Tommy Atkins’, ‘Irwin’ and‘Van Dyke’ mango fruit

Cultivar Disorder first detected (weeks after fruitset)*

Soft noseJelly seed SEC

88 14Tommy Atkins14Irwin 12 12

Van Dyke 88 14

*Fruit were mature 14 weeks after fruit set.


Fig. 2. Photomicrographs of a) cross section (10× ) of the peduncular extension and b) mesocarp section (20× ) of mature fruit withno disorder symptoms. TC, tanniferous cell; RM, resin mass; PA, parenchyma; XY, xylem; RD, resin duct; SC, secretory cell; HWT,helicoidal wall thickening; CW, cell wall; SG, starch grain; FB, fibre bundle.

respectively. Fruit of cv. Irwin did not showsymptoms of these disorders until 12 WAFS,when the fruit were about 68% of their finalweight. Early symptoms of jelly seed and SECwere observed in both immature and mature fruit,whereas soft nose was only observed in fullydeveloped fruit (14 WAF).

3.1. Histology of healthy fruit

In cross-section, tissues of the fruit base hadclosely adhered cells (Fig. 2a), with xylem vesselsintact and undisturbed. The surrounding par-enchyma, including tanniferous cells, or starchidioblasts and resin ducts, also appeared intact,


and it was possible to observe the presence ofsome resin masses. In longitudinal section, thecells were in perfect cohesion, resin ducts withtheir secretory cells and procambial strands wereintact, and intact helicoidal wall thickenings of thexylem elements were also discernible. The meso-carp of a healthy, immature mango fruit was notvery different from that of a healthy, mature fruit.However, more starch grains were observed in thecells of the green fruit mesocarp (Fig. 2b) than inthe cells of the mature fruit mesocarp (Fig. 2b).Fibres with reduced wall thickening were alsoobserved. When the healthy fruit were mature, thecells of the mesocarp appeared empty, i.e. thecellular content was lost but cell cohesion per-sisted.

3.2. De6elopment of stem-end ca6ity

Examination of the early stage of SEC did notreveal significant alterations in the peduncularextension of the fruit. After the transverse cut wasmade, it was possible to observe a series of browndots circularly distributed and located at the open-ings of the resin ducts. The remaining tissuesinside and outside the circle of brown dots ap-peared to be intact. Although the disorder was stillat its early stage, latex exudation was less thanthat observed in healthy fruit, and the exudedlatex had a darker color than that of healthy fruit,which was of milky appearance in both immatureand mature fruit.

In cross-section of the peduncular extension ofthe fruit, tissues appeared partially disrupted. Thedisorganization mainly affected the parenchymacells and the resin ducts in which the secretorycells had disintegrated. Little damage was appar-ent in the xylem, but in some vascular bundlessome tracheary elements were separated from theneighbouring elements of the bundles. In the areaswhere the wall of the parenchyma cells were bro-ken, the cellular contents were lost or dispersedand free tannin granules and starch grains wereeasily observed (Fig. 3a). More Ca oxalate crystalswere observed in sections made from tissues withSEC than in sections from fruit without the disor-der. In longitudinal section, areas of intact cellsalternated with areas of disintegrated cells and

broken cell walls. In intact areas, the cell contourswere clearly defined and the cellular contents werepreserved. In areas where the cells were destroyed,the vascular tissues were altered, and the secretorycells were absent from the inner layer of the resinducts (Fig. 3b).

At the intermediate stage of SEC, there was abrown discoloration at the center of the peduncu-lar extension of the fruit; surrounding areas wereoften still green, and the flow of latex was greatlyreduced. A cavity had formed at the proximal endof the fruit between the skin and the stone, result-ing from the destruction of the vascular connec-tions that linked the seed to the mother plant. Thetissues around the cavity were necrotic and dry,and the inner mesocarp had a yellow coloration incontrast to the outer mesocarp that remainedgreen. At the cellular level, the damage to theparenchyma and to the vascular tissue appeared tobe more extensive at the intermediate stage com-pared to the early stage. In cross-section, largeportions of the parenchyma cells and the tannifer-ous cells had coalesced, and resin ducts wereindistinguishable. Few starch grains were present,and a portion of the xylem from the vascularbundles was broken down (Fig. 4a). In longitudi-nal section, a high degree of tissue breakdown wasobserved, resulting in cell wall fragments and freestarch grains interspersed with intact cells. Theprocambium was partially destroyed and Ca ox-alate crystals were also present. The presence ofthe calcium crystals and starch grains was verifiedwith light polarizing filters (Fig. 4b).

When SEC reached the advanced stage, thecavity was considerably enlarged further. The seedwas totally disconnected from the peduncle, andnecrosis had reached the surrounding mesocarp atthe proximal end of the fruit. The interior of themesocarp had a darker yellow coloration, but nobreakdown was observed in the interior of themesocarp around the stone. In cross-section, mostof the starch idioblasts were destroyed; a fewtanniferous cells and tracheary elements remained.No resin ducts were distinguishable and moststarch grains had disappeared (Fig. 5a). The longi-tudinal section also showed a breakdown of thevascular tissues, the xylary elements had deterio-rated and the procambium was destroyed (Fig.5b).


Fig. 3. Photomicrographs of a) cross section (20× ), and b) longitudinal section (10× ) of the peduncular extension of a mango fruitat an early stage of development of stem-end cavity. XY, xylem; DA, disrupted area; CR, crystal; TC, tanniferous cell; RD, resinduct.

3.3. De6elopment of jelly seed

When fruit with symptoms of jelly seed werecut open, the interior of the mesocarp, i.e. theportion of the mesocarp located next to the seed,was yellow, which contrasted with the surround-ing greenish exterior mesocarp. The latex flowed

normally from the proximal end of the fruitthrough the resin ducts. Most cells of the symp-tomatic mesocarp appeared intact. The number ofstarch grains present in the cells was lower thanobserved in the healthy mesocarp. In general, cellcohesion was maintained and the organization ofthe tissue was preserved (Fig. 6a). At the interme-


Fig. 4. Photomicrographs of a) cross section (20× ) and b) longitudinal section (10× ) of the peduncular extension of a mango fruitwith stem-end cavity at an intermediate stage of development. DXY, destroyed xylem; IXY, intact xylem; FSG, free starch grain;DPA, destroyed parenchyma; CR, crystal; PCA, procambium.

diate stage of jelly seed, the interior of the meso-carp had an orange coloration localized aroundthe seed, whereas the exterior of the mesocarp wasgreen or pale yellow. The affected portion of themesocarp was softer than the non-affected por-tion, and exuded juice if a slight pressure wasapplied. Microscopic observations revealed exten-sive disintegration of walls of most cells. At the

advanced stage of jelly seed, softening of theinterior of the mesocarp occurred, which had theconsistency of jelly or paste, and a brown col-oration around the stone. The tissue appeared tohave completely deteriorated, the walls of almostall cells were broken, large air spaces were ob-served, and starch grains were interspersed amongthe mass of broken cell walls (Fig. 6b).


Fig. 5. Photomicrographs of a) cross section (20× ) and b) longitudinal section (10× ) of the peduncular extension of a mango fruitwith an advanced stage of stem-end cavity. IXY; intact xylem; DXY; destroyed xylem; DPA; destroyed parenchyma; CR; crystal.

3.4. De6elopment of soft nose

The early symptoms of soft nose were charac-terized by a dark yellow color of the mesocarp atthe distal end of the fruit, between the apex of thefruit and the stone apex. The remaining tissues ofthe fruit were intact. Unlike jelly seed and SEC,the first signs of soft nose were not observed inimmature mangoes. Microscopically, cell cohesion

was observed among most of the cells in thesymptomatic mesocarp. A few cells were de-stroyed, and there were fewer starch grains ineach cell as compared to that in the cells ofhealthy mesocarp. As the disorder progressed tothe intermediate stage, the affected area enlargedand reached the endocarp distal end. The affectedtissue appeared softer and exuded juice with onlyslight pressure. The section of the mesocarp af-


Fig. 6. Photomicrographs of the internal mesocarp of a mango fruit at a) early (10× ) and b) advanced stages (10× ) of jelly seed.SG, starch grain; BCW, broken cell wall; ICW, intact cell wall.

fected by soft nose at the intermediate stage hadextensive cellular breakdown, and only a fewdiscernible starch grains. When soft nose was inan advanced stage, a large portion of the meso-carp was destroyed. The affected area includedthe sinus or beak of the fruit, and had reachedthe concave portion of the stone. Unlike jelly

seed, the breakdown did not entirely surroundthe endocarp. At this stage, the disordered tissuehad the consistency of paste, and was softerthan that of healthy, ripe fruit. The reducedfirmness at the distal end was an accurate, non-destructive indicator of the presence of the dis-order.


4. Discussion

There are macroscopic differences among jellyseed, soft nose, and SEC. Each type of internalbreakdown affects different areas of the fruit. Thesymptoms of SEC as described in the literatureare mostly based on observations of ripe fruit(Galan-Sauco et al., 1984; Winston, 1984). How-ever, in the present study, it was observed that thecavity was formed before the fruit were fullydeveloped. Similarly, Malo and Campbell (1978),indicated that flesh breakdown may start beforefruit are fully mature. A brown coloration wasobserved at the openings of the resin ducts at theearly stage of SEC. Deposits of tannins in theresin ducts and in the xylem elements of fruitaffected by black tip necrosis have been reported(Das-Gupta and Asthana, 1944). In that study,the deposits were found in discontinuous, viscousmasses scattered throughout the duct system. Theresin ducts appeared to contain larger masses oftannins than the xylem vessels (Das-Gupta andAsthana, 1944). There is a possibility that tanninswere involved in the brown coloration of the resinducts of fruit proximal end sections with SEC thatwas observed in the present study. Entire resinducts or xylem vessels could not be individuallyexamined. However, it is possible that the accu-mulation of resin in the ducts and in the vesselscould have reached levels that were toxic to thetissues and caused the necrosis observed in theintermediate and advanced stages of SEC. MoreCa oxalate crystals were observed in cross-sec-tions of the peduncular extensions of fruit withSEC than in healthy fruit. While in the crystallineform, Ca may not be available for biochemicalfunctions. Accumulation of Ca in the crystallineform may induce a localized deficiency of avail-able Ca within the fruit, even though the totalconcentration may be similar.

Jelly seed has been considered to be the ulti-mate stage of SEC (Winston, 1984), and has beenincluded among the symptoms of flesh breakdown(Malo and Campbell, 1978). However, in ourstudy, jelly seed symptoms first appeared at thesame time as SEC symptoms, when the fruit werestill immature.

Soft nose has been described as a combinationof jelly seed and SEC (Galan-Sauco et al., 1984).The present study shows that there are differencesamong SEC, jelly seed, and soft nose. Jelly seedaffected only the interior of the mesocarp,whereas SEC affected the proximal end of thefruit. Soft nose symptoms first appeared when thefruit were close to maturity and, although thedisorder may affect a large portion of the meso-carp in the advanced stage, soft nose mostly af-fected the fruit mesocarp distal end. Thesymptoms of jelly seed developed all around theendocarp, whereas those of SEC may also affectthe interior mesocarp entirely or partially. Stem-end cavity was associated with deterioration ofthe vascular tissues between the stone and themesocarp, and necrosis of the mesocarp aroundthe cavity. However, no cavity, air pocket, gap ortissue necrosis developed in fruit affected withjelly seed or soft nose. Consequently, prematuresenescence of the mesocarp observed in SEC, jellyseed, and soft nose may not be caused by thesame factors.

Although the visual symptoms of the differenttypes of disorders appeared different, no majormicroscopic differences were established amongjelly seed, soft nose, and SEC. In all three disor-ders, disorganization of the cells and rupture ofthe cell walls appeared as the first microscopicindicators. Once the cell walls were disrupted, thecellular contents were released and, as timepassed, the disorders extended to areas that wereintact, until the affected tissues became watery,paste-like, or jelly-like.

It was not possible, using light microscopy, toobserve in detail the alterations caused to the cellwalls. The literature does not contain informationabout the biochemical changes associated withjelly seed, soft nose, or SEC. However, researchon other physiological disorders of mango indi-cates that significant changes occur in the activi-ties of several enzymes present in the mesocarp ofdisordered fruit. The activities of malic enzymeand pectin methylesterase increased in the meso-carp of cv. Alphonso mangoes affected withspongy tissue (Krishnamurthy, 1981). Also, amy-lase and invertase activities were lower in meso-carp with spongy tissue than in healthy pulp


(Katrodia, 1988). However, as observed in fruitwith spongy tissue, the breakdown observed injelly seed, soft nose, and SEC was probably theresult of enzymatic activities.

Tissues with jelly seed and soft nose symp-toms contained fewer fibre strands than healthymesocarp. This may have been caused by a de-struction or dissolution of the fibres early in theonset of the disorders. Similar deterioration ofthe vascular tissues was observed in SEC. Thisbreakdown may have isolated the rest of themesocarp from the seed, interrupting the supplyof nutrients to the mesocarp. Although fruit Cadeficiency early during fruit ontogeny has beenimplicated as the cause of internal breakdown inmango (Burdon et al., 1991), the present studycould not confirm or deny this. It is possiblethat the different internal breakdown symptoms(soft nose, jelly seed, and SEC) have a commoncause and that Ca deficiency is involved. How-ever, based on temporal and spatial differencesin symptom development within the fruit, it ap-pears that soft nose, jelly seed and SEC may beclassified as separate disorders.

Acknowledgements

The authors thank Mr. Frank Sesto for tech-nical assistance, Brooks Tropicals, Inc. for useof their orchards, the Tropical Fruit Growers ofSouth Florida Advisory Council for financialsupport of the project, and Drs. B. Dehgan andE.A. Hanlon for critical review of themanuscript. This article is Florida AgriculturalExperiment Station Journal Series No. R-05845.

References

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