1537 11 012 0.001). In ) produced produces a < P dry weight. Mean However, the levels 1 − -tomatine content (in 10 Halley 3155 α cerasiforme gg -tomatine appears to be µ α var. L. cv. dry weight) and even at this 2012 Society of Chemical Industry 1 c − -tomatine content in tomato. ∗ α a 0.001) different between cropping years, -tomatine is poorly absorbed in the gut and < α P species, is a steroidal glycoalkaloid which functions Lycopersicon esculentum -Tomatine has been reported to have numerous health α Lycopersicon esculentum -tomatine content in tomatoes. -tomatine content than their conventional counterpart α α Department of Plant Sciences,Davis, One Davis, CA, Shields 95616, USA Avenue, University of California Correspondence to: AlysonTechnology, E One Shields Mitchell,95616, Avenue, Department USA. E-mail: University [email protected] of of Food California Science Davis, Davis, and Department CA of Food Science andof Technology, California One Davis, Shields Davis, Avenue, CA, University 95616, USA In general, most glycoalkaloids are considered to be a negative Solanum ∗ a b attribute due to their inherent toxicity. red ripe tomatothe fruit range with of a 500–5000 very mg high kg is hydrolyzed to thestudies relatively of glycoalkaloids non-toxic over aglycone. thepossess More past beneficial decade recent effects suggest depending thattheir on they use. dose and conditions of It appears that level, human consumption of this fruit produces no ill effects. of glycoalkaloids consumedconsidered to from be red–ripe toxic tovariant humans. tomatoes of In fact, are in Peru not an indigenous -tomatine content in tomatoes was also observed and suggests α and some -type -tomatine ) is found α and Alyson E Mitchell 21 Solanum -tomatine and b NO α 83 Levels are highest Lycopersicon alkaloids have one H fresh weight. As the 2 50 1 Glycoalkaloids are not − -tomatine content was also significantly ( 1,2 α Production of Solanum The levels of 6,7 -type alkaloids). The aglycone 9 species. -tomatine (C α -Tomatine has numerous potential health benefits including the ability to inhibit cancer cell growth The primary glycoalkaloids present α 5

Solanum – 3 -tomatine were compared in dried tomato samples ( Stephen Kaffka : 1537–1542 www.soci.org α -tomatine; organic; conventional -Tomatine is influenced by numerous agronomic factors including fertilization and nitrogen availability. Solanum -tomatine and dehydrotomatine (Fig. 1). a Because of these high levels, synthesis is α α 93 α

-Tomatine, synthesized by fresh weight, respectively. 8 -tomatine levels of tomatoes in both cropping systems ranged from 4.29 to 111.85 α and 1 α − 2013; -tomatine were significantly higher in the organically grown tomatoes than conventional ones ( tomato; studies. α

-tomatine in tomatoes Lycopersicon 2012 Society of Chemical Industry in vitro c dehydrotomatine in tomatoes ranged from 521to to 1498 16 mg kg 285 and 42 J Sci Food Agric The steroidal glycoalkaloid INTRODUCTION BACKGROUND: α Abstract practices on the content of the glycoalkaloid Eunmi Koh, organic and conventional crop management (wileyonlinelibrary.com) DOI 10.1002/jsfa.5951 A long-term comparison of the influence of Research Article Received: 3 August 2012 Revised: 25 September 2012 Accepted article published: 15 October 2012 Published online in Wiley Online Library: 9 November 2 tomato ripens and changes colorthe from glycoalkaloids green decrease. to red, the levels of glycoalkaloids is favored bythe development the of chlorophyll. same The concentration conditions of is that highest promote inhigh stems, levels leaves, (up to and 5%fruits green fresh of weight) tomatoes. in tomato. It leaves, flowers can and reach green portion is called tomatidine. These These glycoalkaloids are ethersa between carbohydrate a moiety steroid ( alkaloid and nitrogen atom located in the F ring. in tomatoes are sensitive to nitrogen availability. in required for plant growthin and function. plant However, they defense function and mechanisms predatory against insects. fungi, bacteria, viruses in immature green fruit up to 500 mg kg to protect against pathogens and insects. Although glycoalkaloids are generally considered toxic, levels of suggesting that other influencing factors such as environmental conditions also affect well tolerated in humans. the organic management system, CONCLUSIONS: The organically produced tomatoes had higher average in Keywords: over the 10-year study. Significantthat environmental annual factors, variability external in to nitrogen the fertilization, influence in organic and conventional croppingResearch systems on that Agricultural had Systems been project archived (LTRAS) over at UC the Davis. period fromRESULTS: 1994 to The 2004 from the Long Term Herein, the levels of 23 . 3 et al. 3 et al CH CH : 1537–1542 93

N H

N 2013; H O O C 3 C J Sci Food Agric 3 H 3 H 3 -tomatine dehydrotomatine CH α CH 3 3 CH CH Both conventional and organic agricultural practices include affect the nutritive compositionplant metabolites. of Organic plants, systems emphasizeof including the soil secondary accumulation organic mattercover and crops, fertility manures over and time compostsdiverse through and soil rely the ecosystem on use to the of makeavailable activity to nitrogen plants. of Conventional (N) farms a utilize and fertilizers containing othersoluble nutrients inorganic nitrogen anddirectly available other to plants. nutrients, The availability which of inorganic are nitrogen,particular, in has the more potential to influence the synthesis ofplant secondary metabolites, proteins and soluble solids. Mitchell proposed that increased tomatorates, crop and growth greater and development biomasswould accumulation in also well-fertilized correlatetowards crops the with production decreasedcontaining of secondary allocation starch, metabolites. of celluloseplant Given metabolites and that resources are many non-nitrogen producedand secondary are for inducible by pathogens defense or wounding, againstin possible pest differences herbivores pressure between conventional and organic systemsalso might influence levels in food crops. combinations of farming practicesupon that region, vary climate, greatlyfactors soils, depending guiding pests the and particular diseases, management and practices economic used on D-galactose O D-galactose O OH OH In a O www.soci.org E Koh O 14 OH HO OH HO 2012 Society of Chemical Industry -tomatine in O Studies also α c O 20 O O – O O 9,18 OH -tomatine is a secondary OH OH α OH O O D-glucose D-glucose D-glucose O D-glucose O OH OH HO HO -tomatine content of tomatoes is α HO showed a negative correlation between HO and enhancing the immune system. OH 15 . OH 13 HO -tomatine and dehydrotomatine. -tomatine has high activity against prostate HO α α et al 16,22,23 The influence of different types of fertilizers and 16,17 2,21 lowering plasma low-density lipoprotein cholesterol and that 12 Structure of HO HO xylose HO xylose HO Fundamental differences between organic and conventional Multiple factors including genetics, soil type, fertilization in vitro -tomatine and macrophage expression of tumor necrosis factor- and triglyceride levels, production systems, particularly in soil fertility management, may practices, ripening stage,influence and abiotic glycoalkaloid and/or biotic levels stress in can plants. cropping system on the production ofbeen bioactives investigated. in tomatoes has cancer cells. Given the data in support of a role for α α the prevention of these chronic diseases, interest in understanding the levels in tomatoes has increased. As plant metabolite, similar tolevels may other be influenced secondary by the plant agronomicthey environment are metabolites, in grown. which wileyonlinelibrary.com/jsfa Figure 1. promoting properties including: anti-cancer activity against tumor cell lines, demonstrate thatdependent the on thetomato tomato plant. variety and anatomical part of the recent study, Choi

1538 1539 - 10 α . Vicia = 3) for et al n of ammo- 1 (10 000 per − 1 − year in addition 1 − L.). Transplanting of all sub-sample areas (sub- -solanine was added as an 2 α wileyonlinelibrary.com/jsfa of composted manure was added to Pisum sativum 1 − C, ground, and stored in glass containers ◦ C until analyzed. Samples from conventional standard solution of ◦ 1 − -tomatine -tomatine analysis. Three of the sub-plots ( α α of an N-P-K starter fertilizer and 118 kg ha of composted poultry manure currently is top dressed 1 1 − − Roth) and field peas ( Crop management practices follow best management practice Lofa1mgmL -Tomatine analysis followed the method of Cataldi total biomass were measured everyand year and C. analyzed Sample for archiving totalfrom included N all yearly cropping systems, plant and time and zero andples fruit subsequent soil collected samples sam- every few years.were Systems compared, rather so a than valid single comparisonwas inputs managed required carefully that to achieve each its system potential yields.both For the example, conventional and organic maize/tomato systems hadsame the tomato cultivar (Halley 3155) over thewere period irrigated of similarly interest and as neededof but organic N the sources short-term can availability requireN more needs. total The N input systems toinclude compared meet representative crop crops are rather model thanable systems, more complex, crop chosen change- to rotations.rotations California as markets farmers change, and organic rarely farmers,to especially, have follow tend more complex fixed rotations than crop the one studied at LTRAS. guidelines in50 kg the ha region. Conventional tomatoes received internal standard. Internal standard recovery was 92–93% for solanine. The mixture was extracted at room temperature for 1 h by A 250 mgcombined of with 5 mL pestle-pulverized, of 1% acetic air-driedµ acid in water. tomato To this sample mixture, 25 was tomato crops. This wasincreased reduced to after a organic near matterAugust constant each levels year level. when had the Tomatoes field as were aA whole commercial harvested reached tomato 90 in % harvester ripe was fruit. used for main plot harvests. Sampling and preparation of plant material Immediately prior to harvestarchive of were the collected from main four plot, 3.1 samples m for the plots) within the largeryield main of plots. the Total red plantrandom ripe sample biomass of fruit 20 and ripe fruit were the from the determinedand four sub-plots for oven was washed each dried sub-plot. at A 60 and incorporated with the coverin cover crop. crops The varies from amount year ofcurrently to receive N year, between but present 240 typically, and organic 260 plots kg N ha nium nitrate at side-dress. A combination ofwere tillage used and herbicides for weedperiodically control. and Aphids, were controlled mites as and necessary,in similar stinkbugs commercial to occur practices fields.were In transplanted both at treatments, the rate processing of tomatoes 22 500 plants ha and organic plots from even-numberedchosen years for (1994–2004) were each treatment (three organic; three conventional) werein analyzed each of thewere even analyzed in years triplicate. (1994–2004). These sub-plot samples Analyais of in the dark at 20 villosa acre). Transplanting in the organic treatmenttion followed of incorpora- a winter legume cover crop consisting of hairy vetchplots ( occurred within aaround period the middle of of April. 2–4 Prior to days incorporation of9Mgha in the cover spring, crop, commonly to the N fixedcropping by cycles, to the more rapidly legume increase soil cover organicand matter soil crop. levels fertility, During 19 Mg the ha first three α ) 4)- → However, -tomatine α 26 2012 Society of Chemical Industry Halley 3155 – c 22 -glucopyranosyl- Sorghum vulgare D L. cv. - β -glucopyranosyl-(1 D - β -tomatine in tomatoes www.soci.org α 3)]- → )-spirosolan-3-yl cartridge (1 g, 6 mL), acetic acid (glacial), S 18 ,25 : 1537–1542 Lycopersicon esculentum α -Solanine (99%) was obtained from Sigma (St 93 ,5 α β 2013; -xylopyranosyl-(1 D - β 2)-[ -galactopyranoside, was purchased from MP Biomedicals Inc. The present study is an example of the use of long-term research → D - -Tomatine, (3 (Solon, OH, USA). Louis, MO, USA). The C L.) in 1992established in and 1993 1993,was using replicated three ten 0.4-ha times and plots. different bothwere crops Each present of cropping the cropping each 2-year systems rotations the year. system use were Plots of commercial were scalewere large farm measured equipment. enough using Irrigation flow to amounts metersSystems allow located differ at for in each the irrigatedirrigated), plot. in amount the of amounts of irrigation nutrientsorganic received matter applied applied as (rain to fertilizers the fed and soil in cover as or crops, crop and/or residues, composted winter manure (http://asi.ucdavis.edu/rr). legume Conventional plots received herbicidesneeded and while organic other crops pesticidespesticides received as such only as organically sulfur approved made and Bt by compounds the (a bacterium natural Bacillus pesticide thuringiensis). Crop yields and Tomato cultivation in the LTRAS cropping systems Following uniform cropping with Sudan grass ( and all solvents were from Fisher Scientific (Fair Lawn, NJ, USA). J Sci Food Agric MATERIALS AND METHODS Chemicals and samples α Cropping system comparisons of the farm. Manya of steady thesefarms. state influences The condition change dynamicadequately may continuously, nature controlled comparisons of so never of agricultural produceconfounding quality, be systems free also from influences, achieved makes of experimentally on studies challenging. most comparingand Reviews the organically nutritional produced quality vegetablesdifferences of demonstrate with the inconsistent conventionally exception(vitamin of C) higher and levels less of nitrate ascorbic in acid organic products. (1 to address complexThe processes specific operating objective in was to cropping compare systems. the levels of these data are difficultagronomic to conditions interpret varied sincesampling widely cultivar and and selection and analysis differentIn methods were contrast, of comparisons used ofresearch in plots cropping the that systems, have using investigationsprovide been a long-term cited. managed means consistently toassociated over overcome with time, many farm-based of sampling. the Additionally,changes confounding the over effects factors time of in cropping systemusing behavior archived can soil be and evaluated plantof samples the and causes of a those reasonable changes estimate can be made. in tomato samplesproduced ( in conventionalhad and been organic archived croppingLong over Term systems the Research that period onUC from Agricultural Davis 1994–2004 Systems which project from began (LTRAS)was designed the in at to detect 1993 and estimate (http://ltras.ucdavis.edu). changestrends in and LTRAS crop other factors productivity correlated with sustainability, which result from differing irrigationan and organic fertilization cropping systemgrown practices. in in It which rotation includes maize andconventionally. and compared This tomatoes to archive are theCalifornia and same perhaps of the crops world. tomato produced samples is unique in β - α 27,29 et al. To date, 9 . : 1537–1542 dry weight) in etal 93 1 -tomatine. In the − α 2013; gg µ demonstrated that the 9 . et al reported that total glycoalkaloid -tomatine content was observed -tomatine levels are significantly J Sci Food Agric α 31 α -tomatine levels in the tomatoes pro- -tomatine content is only statistically α -tomatine content ( α found that increases in the glycoalkaloid α It appears that this influenced the levels 32 . -Tomatine, however, is primarily a carbon- α etal 29,30 27,28 -tomatine levels were higher in the organic plots. α Comparison of dry weight) in the organic (A) and conventional tomatoes 1 − 0.001)differentamongcroppingyears(Fig.3A),whereasinthe -tomatine in these tomatoes; the 1998 tomatoes receiving α Figure 3 demonstrates the changes in the tomatine content < gg -tomatine. Mondy and Munshi -tomatine content of tomato leaves correlates with the ratio of µ P conventional system the different in 1996 and 2002 (Fig. 3B). Totaluouslydecreasedfrom1994to1998intheorganicsystemandthen N application rate contin- increased almost up to the starting level intional 2000, while system the received conven- the same totalIn N both input cropping during systems, 1994–2004. duced in 1996 were highest, although not statisticallyconventional higher in system the (Fig. 3A and B). Theorganic plots soil at organic LTRAS reached matter a in quantitative limit the of accumulation in 1998 and the amount ofplots composted manure was applied to reduced. organic ( organically and conventionally grown tomatoes from 1994with to same 2004. letters Means are not significantly different. (B) collected over 10 years oforganic the cropping LTRAS system, trials (1994–2004). the ( In the based compound (Fig. 1). Hoffland In organic plots, the soilbefore microbes it can must be assimilated. first This release can result theduction in of a nitrogen more primary balanced and pro- secondary plant metabolitesexplain and why may help Figure 2. content (e.g.readily proteins, available. DNA, peptides, etc.) when soil N is of the lowest nutrient applicationα had significantly lower levels of studies comparing the phytochemicalsand content organic of crops conventional demonstratepart, inconsistent can differences. be This, explained by in thefrom relative various rates fertilizers, of release which ofplants, can and nutrients influence the result production of in secondary metabolites. various C/N ratios in content of potatoes increased significantlyof with N increasing fertilization levels when ammonium nitrateAdditionally, Cronk was applied to the soil. α C/N and proposed that theN carbohydrate concentration, moiety, limits rather the than biosynthetic the ratepresent of study, a higher average intheorganictomatoesascomparedtotheconventionaltomatoes (Fig. 2). This would suggest that the readily available nitrogenconventional in plots the is not a primary factor in the biosynthesis of tomatine and supports the observations by Hoffland m µ www.soci.org E Koh -tomatine -tomatine. α α 2012 Society of Chemical Industry c -Tomatine was 2.00 mm (Phe- α × ) ion source. Sam- plants will empha- who found that the 27 dry weight) most years. 20 1 ZSPRAY -solanine, was determined − α ), 150 mm -tomatine and solanine was µ gg α µ 0.001) higher in the organically dry weight) as compared to the wilt broke out in 2004 in both Tomatosamplesinthisstudywere . The cone voltage (20–89 V) and 1 1 -tomatine content in ripe tomatoes. < 21 − − α P used for the solid-phase extraction). C, respectively. Peaks corresponding 0.05 was considered significant. ◦ gg 18 < µ Verticillium value P C and 400 on a dry weight basis. This is similar to results ◦ 1 − fresh weight when considering that the moisture content gg 1 µ -tomatine levels of tomatoes ranged from 4.29 to − α -tomatine and solanine were identified by full scanning liq- -tomatine to an internal standard, SPE cartridge (1 g, 6 mL, Thermoscientific, Bellefonte, PA, USA), gg α α According to the C/N balance hypothesis, Tomatine was resolved by reversed-phase high-performance µ -tomatine content of ripe red tomatoes ranged from 0.3 to -tomatine levels was observed between two different cropping 18 nomenex, Torrance, CA, USA).with The a HPLC SIL-10A systemdetector injector, was (Shimadzu binary equipped Scientific). 10 AD0.1% The pumps, mobile formic and phase acid SPD-10A gradientacetonitrile in UV was water (solvent (solvent B):15 A) 35–38% min, and 38–65% B 0.1% B10 in formic in min 5 acid 5 min, between min. in The 38% runs.LC column triple-quadrupole B The was mass spectrometer from re-equilibrated HPLC (Micromass, for UK) 5 Altrincham, was equipped to with interfaced a to dual orthogonal a ( Quattro which was preconditioned withwater, 6 passed mL through of C methanol and 6 mL of offreshtomatoisaround95%. collision energy (55–80compound V) for were accurate quantitative optimized data.of separately The ratio for of each peak area and then plotted as a function of the concentration of Statistical analysis The data weremultiple analyzed range tests statistically using the byCary, SAS NC, ANOVA USA). 9.2 A version and (SAS Institute Duncan’s Inc., wileyonlinelibrary.com/jsfa The 111.85 RESULTS AND DISCUSSION using a Lab-Line Orbit Environ-ShakerMelrosePark,IL,USA).Thehomogenatewascentrifugedfor15 (Lab-Line Instruments Inc, The min. pellet was suspended in 5 mL of 1%andcentrifuged.SupernatantcombinedwasappliedtoaHyperSep acetic acid in water, shaken C During this study, size the synthesis of growth-related compounds with high N ples were run in3.0 kV. positive The ion source mode temperatureture using and were a a 150 capillary desolvation voltage gas of tempera- liquid chromatography (HPLC;MD, Shimadzu USA) using Scientific, a Columbia, Prodigy ODS (5 The sample was washed through using 5% methanoland in eluted water (v/v) with methanol. EluentPTFE was filter filtered (Millipore, through Bedford, MA, a USA). 0.45 to uid chromatograph–electrospray ionization/mass(LC-ESI/MS). spectrometry Structural verificationmolecular of ions corresponding peaks to having ions with achieved by product ion scanning LC-ESI/MS/MS. quantified through multiple reactionlimit monitoring of mode with detection the of 5 ng mL reported earlier by Friedman and Levin, conventionally grown ones (23.16 harvested in the red–ripe stage. The mean levels of (Fig. 2) were significantly ( grown tomatoes (42.25 α 6 systems. The conventional system received treatmentwhereas for the the wilt, organic system didα not. No significant differencesystems in in 2004. This suggests thata pathogen critical stress role may of not determining have

1540 1541 : - ., ., ., α ., et al et al et al J Agric et al Spinacia Plant Soil . cerasiforme :3891–3899 J Food Sci -tomatine and J Food Compos 60 Phytochemistry δ var. :357–368 (1983). :401–403 (1984). J Agric Food Chem :3786–3791 (2009). 40 :143–148 (1977). 77 -, and 57 51 :141–156 (1976). γ Choristoneura-fumiferana Oikos -Tomatine determination Proc Nat Acad Sci USA 61 Botrytis cinerea -tomatine. 1-, α α -tomatine-to-esculeoside A β α -, :8655–8681 (2006). wileyonlinelibrary.com/jsfa α :3217–3222 (2007). Proceedings of the First Scientific 54 Angew Bot :3144–3150 (2012). J Agric Food Chem Rapid Commun Mass Spectrom :6154–6159 (2007). 581 60 :897–900 (2000). 55 65 J Agric Food Chem -Tomatine content of tomato and tomato Am J Potato Res Lycopersicon esculentum :279–294 (2007). Ann Entomol Soc Am α FEBS Lett :1507–1511 (1995). :5751–5780 (2002). 744 . 43 50 J Food Sci J Agric Food Chem :1959–1964 (1994). J Agric Food Chem 42 :140–152 (1999). J Agric Food Chem Acta Hortic L.). 18 :20–27 (2006). 19 :263–272 (1999). :441–451 (2007). :9–25 (1974). :3103–3110 (2005). :12877–12881 (1994). :2832–2839 (2004). feeding deterrents for spruceLepidoptera budworm Tortricidae. Tomatidine, the major saponin in tomato, induces programmeddeath cell mediated by reactive oxygen speciesFusarium in oxysporum the fungal pathogen (2012). Gruppen H, C22conversion during tomato isomerization ripening is in of driven triterpenoidal by skeleton. C27 hydroxylation on the major antioxidant componentsAnal of tomatoes. of potato glycoalkaloids. different stages of development. products determined by HPLC with pulsed amperometric detection. J Agric Food Chem in tomatoes by HPLC using pulsedFood Chem amperometric detection. differences inorganic processing tomatoes and72 grown conventional under production commercial systems. Ten-year comparison of the influencecrop of organic and management conventional tomatoes. practices on the contentqualities. of flavonoids in J Agric Food Chem and in the diet. 210 Structure–activity relationships of tomatidine against human breast (MDA-MB-231), gastricand (KATO-III), prostate (PC3) cancer cells. conventional croppingflavonoids, systems nitrate, and oxalate on in 27 ascorbic varieties of spinach acid, ( vitamin C, oleracea plants in relation to vertebrate herbivory. conventional maize andterm tomato experiment cropping in systems California, from in a long- 13 and susceptibility of tomato leaves to by liquid chromatographytandem coupled with mass electrospray19 spectrometry. ionization in ripe fruit of Andean developmental and91 genetic aspects. Glycoalkaloids andcolon metabolites (HT29) inhibit and the liver (HepG2) growth cancer of cells. human 52 hamsters reduces their plasma low-densityand lipoprotein triglycerides. cholesterol aggregate structure adjuvants modulate lymphocyteand proliferation Th1 and Th2 cytokineVaccine profiles in ovalbumin immunized mice. 4 Bentley MD, Leonard DE and Bushway RJ, Solanum alkaloids as larval 5 Ito S, Ihara T, Tamura H, Ikeda T, Kajihara H, Dissanayake C, 6 Friedman M, Tomato glycoalkaloids: Role in the7 plant and in Friedman the M, diet. Potato glycoalkaloids and metabolites: roles in the plant 8 Roddick JG, The steroidal glycoalkaloid 9 Hoffland E, van Beusichem ML and Jeger MJ, Nitrogen availability 16 Yamanaka T, Vincken JP, Zuilhof H,17 Legger ToorRK,SavageGPandHeebA,Influenceofdifferenttypesoffertilizers A, Takada N and 18 SindenSL,SanfordLLandWebbRE,Geneticandenvironmentalcontrol 19 Ali A and Schloesser E, Tomatine content of plant20 parts of tomatoes Friedman at M and Levin CE, 21 Friedman M, Levin CE and McDonald GM, 22 Barrett DM, Diaz WJV and Watnik M, Qualitative23 and Mitchell nutritional AE, Hong YJ, Koh E, Barrett DM, Bryant DE, Denison RF, 24 Dorais M, Effect of cultural management on tomato fruit health 15 Choi SH, Ahn J-B, Kozukue N, Kim H-J, Nishitani Y, Zhang L, 25 Koh E, Charoenprasert S and Mitchell AE, Effect of organic and 26 Bryant JP, Chapin FS and Klein DR,27 Carbon/nutrient balance of Kaffka boreal SR, Bryant DC and Denison RF, Comparison of organic and 10 Cataldi TRI, Lelario F and Bufo SA, Analysis of tomato glycoalkaloids 11 Rick CM, Uhlig JW and Jones AD, High alpha tomatine content 12 Lee KR, Kozukue N, Han JS, Park JH, Chang EY, Baek EJ, 13 FriedmanM,FitchTE,LevinCEandYokoyamaWH,Feedingtomatoesto 14 Rajananthanan P, Attard GS, Sheikh NA and Morrow WJW, Novel J JSci to solanine 2012 Society of Chemical Industry -carotene and β c dry weight) in organic (A) The enzymes that are 1 − 33 Beauveria bassiana gg 0.05. µ = -tomatine in tomatoes www.soci.org α α -tomatine content in tomatoes. α :2079–2083 (2004). : 1537–1542 52 -tomatine content ( 93 :697–706 (1989). α :195–200 (2003). 15 83 2013; -tomatine content than their conventional counterpart α Changes of -tomatine content in tomato fruits and vegetable plant tissues. α lycopene content in tomatoes duringFood growth Agric and maturation. and tomatine: Plant defensive chemicals inhibit an insectJChemEcol pathogen. Agric Food Chem -tomatine content in tomatoes treated with the same N α In conclusion, the organically produced tomatoes had higher (B) (A) 2 Kozukue N, Han JS, Lee KR and Friedman M, Dehydrotomatine and 3 Costa SD and Gaugler RR, Sensitivity of 1 Kozukue N and Friedman M, Tomatine, chlorophyll, and conventional tomato (B) over 10Within years of the the LTRAS cropping trialletter system, (1994–2004). are tomatine not significantly content different at annotated by the same J Sci Food Agric REFERENCES content due to N fertilizationvariety varied comparisons with have potato not been cultivar.in made Similar N in tomatoes. fertilization Increases content have of been the shown plant to through increased. increase the chloroplast Figure 3. Cropping system comparisons of application over 10 years (conventional plots)This was suggests also observed. thatfertilization, environmental influence affect factors, external to nitrogen when using thethe same tomato cultivar. 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