Bt Brinjal in Bangladesh: The First GeneticallyEngineered Food Crop in a Developing Country

Anthony M. Shelton,1 Md. J. Hossain,2 Vijay Paranjape,3 Md. Z.H. Prodhan,4 Abul K. Azad,4

Ritu Majumder,3 Sayed H. Sarwer,2 and Md. A. Hossain2

1Department of Entomology, Cornell/NYSAES, Geneva, New York, 14456, USA2Feed the Future South Asia Eggplant Improvement Partnership, Ithaca, New York 14853, USA3Sathguru Management Consultants Private Limited, Banjara Hills, Hyderabad 500034, India4Bangladesh Agricultural Research Institute, Gazipur 1701, Bangladesh

Correspondence: ams5@cornell.edu

Eggplant, or brinjal (Solanum melongena), is a popularly consumed vegetable grownthroughout Asia that is prone to vicious and sustained attack by the eggplant fruit andshoot borer (EFSB) (Leucinodes orbonalis) throughout the growing season. Yield losses inBangladesh because of EFSB infestation have been reported as high as 86%. Farmersreduce crop losses by frequent applications of insecticide. To counter the EFSB pest,Bangladesh has developed and released four Bt brinjal varieties expressing Cry1Ac(Bt brinjal). Bangladesh is the first developing country to release a commercial geneticallyengineered (GE) food crop. In this article, we discuss the development and adoption of Btbrinjal in Bangladesh from initial distribution to 20 farmers in 2014 to cultivation by morethan 27,000 farmers in 2018. Bt brinjal provides essentially complete control of EFSB, dra-matically reduces insecticide sprays, provides a sixfold increase in grower profit, and does notaffect nontarget arthropod biodiversity. A major focus is to ensure its durability throughstewardship. Bangladesh has shown great leadership in adopting biotechnology for thebenefit of its farmers and serves as an example for other countries.

HISTORY OF Bt CROPS

One of the earliest published reports of agenetically engineered (GE) plant express-

ing insecticidal crystal (Cry) proteins from thebacterium, Bacillus thuringiensis (Bt), was byMonsanto scientists who transformed the toma-to in 1987 (Fischhoff et al. 1987). Their reportwas prescient when the authors wrote, “Theseengineered tomato plants represent a significantstep to increased selectivity, specificity and effi-

cacy in insect control.” In fact, Bt plants haverevolutionized insect pest management in cornand cotton worldwide (Shelton et al. 2002; Ro-meis et al. 2008) by effectively controlling keyinsect pests while conserving their natural ene-mies (Romeis et al. 2019). However, perhaps theauthors could not fully anticipate the extent ofthe social and political activism that their newtechnology would cause.

From an agronomic standpoint, Bt cropsshould be considered another form of a pest-

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resistant crop (Kennedy 2008). They share thesame potential benefits (e.g., reduced need forinsecticides) and liabilities (e.g., insects develop-ing resistance to the Bt proteins they express) asconventionally bred crops with host-plant-resistant characteristics. There is a long historyof conventional breeding to develop crops resis-tant to insect pests and much of this work con-tinues to this day. Breeding for insect resistancehas been especially successful in the case of fieldcrops (e.g., wheat lines resistant to the Hessianfly and alfalfa lines resistant to the soybeanaphid) and many of the genes have been identi-fied (Smith and Clement 2012). However, therehas been little success in developing resistantlines to insect pests in the orders of Lepidoptera(caterpillars) andColeoptera (beetles); these twoorders contain many of the world’s most dam-aging pests.

Host plant resistance is considered a corner-stone of integrated pest management (IPM) be-cause it provides an effective and economic wayby which crop damage can be avoided (Sternet al. 1959; Naranjo and Ellsworth 2009). Btcrops are the first of what can be consideredGE insect-protected crops, but they will certain-ly not be the last.

Bt is a family of bacteria that contains insec-ticidal proteins that, when ingested by certaininsects, cause holes in their midgut that disruptits function and lead to their death (Heckel2012). Bt has been used as a foliar insecticidefor decades and, although it had relatively lim-ited use, it has a stellar safety record for humansand the environment (Shelton et al. 2002). Bthas critical limitations when used as a foliarlyapplied insecticide. First, its short persistence(2–4 days) may lead to frequent applications,which costs the farmer not only money for theproduct but also for its application. Second, be-cause it needs to be ingested by the targetedinsect, thorough coverage of plant parts is re-quired. Such coverage is extremely difficultto obtain in many crops, especially those inwhich the insect bores into tissues, such as theEuropean corn borer that bores into the stalk aswell as into the ear. These limitations were over-come when the genes producing the Cryproteins were introduced into plants and the

proteins were expressed in insect-susceptibleplant parts.

The first GE crop to be commercializedanywhere was in 1995 using potato cultivars ex-pressing the Bacillus thuringiensis var tenebrio-nis Cry 3A toxin against the Colorado potatobeetle (CPB), Leptinotarsa decem lineata (Say)(cv NewLeaf, Monsanto). Bt potato controlledCPBverywell evenunder intensepressure.How-ever, Nature Mark, which marketed the productfor Monsanto, dissolved after the 2001 season(Shelton 2012). The product was doomed be-cause of several factors,most notably the contro-versy about a GE food crop. Such controversycontinues forother fresh food crops.Meanwhile,Bt cotton andmaize have been far more success-ful. In 2017, 59.7millionhectares ofBtmaize and24.1 million hectares of Bt cotton were grownglobally (James 2018).

Bt crops have revolutionized insect pestmanagement by providing host plant resistanceto key pests in the insect orders, Lepidopteraand Coleoptera. Prior to the development of Btcrops, there were few examples of strong hostplant resistance to these insect orders. For de-cades, Bt had limited use as a foliar insecticideprimarily for organic growers, who had limitedalternatives, and for arboreal pests where ecolog-ical sensitivity was critical. However, when in-secticidal genes were incorporated into Bt crops,Bt quickly became a major insecticide. Theadoption rates for Bt crops have been unprece-dented in agriculture (James 2000). Between1996 and 2015, this adoption has been associat-ed with increases in farm income of more than$50,274M and $45,958M in Bt cotton andmaize, respectively, and reductions of morethan 268M and 87M kg of insecticide-active in-gredient in Bt cotton and maize, respectively(Brookes and Barfoot 2017). However, the po-tential benefits provided by Bt crops havelargely gone unrealized in fruits and vegetableswhere insect management continues to relyprimarily on the use of synthetic insecticides.Except for the use of Bt sweet corn, which hasprovided excellent control of ear-infesting cat-erpillars (Shelton et al. 2013), currently no Btvegetables or fruits are commercialized. This ischanging in at least one country of Southeast

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Asia—Bangladesh—with its 2014 release of Bteggplant.

EGGPLANT IN SOUTHEAST ASIA

Solanum melongena L. (eggplant, also known asbrinjal in India and Bangladesh, and talong inthe Philippines) is one of the most important,inexpensive, and popular vegetable crops grownand consumed in Asia. Although its popularityvaries by country in Asia, it is generally con-sidered as one of the top five vegetables inAsia. The biggest constraint to eggplant produc-tion throughout Asia is a chronic and wide-spread infestation by the eggplant fruit andshoot borer (EFSB), Leucinodes orbonalis (Gue-née) (Ali et al. 1980). Caterpillars damage egg-plant by boring into the petiole and midrib ofleaves and tender shoots, resulting inwilting anddesiccation of stems. Larvae also feed on flowers,which results in flower drop ormisshapen fruits.The most serious economic damage caused byEFSB is to the fruit, because the holes, feedingtunnels, and frass (larval excrement) make thefruit unmarketable and unfit for human con-sumption (Fig. 1). To control this insect, farmersroutinely spray broad-spectrum insecticides, of-ten 2–3 times per week, and, in some cases, twicea day (for an example of commonly used insec-ticides, seeDel Prado-Lu 2015). Consequently, itis not uncommon to apply over 100 sprays perseason, resulting in high residues on marketablefruit. Such an insecticide-dependent strategyposes both environmental and health concerns.

Environmental concerns include killing naturalenemies that can help reduce pest populations,leaching of active pesticide ingredients into thesoil and water, and harming pollinators. Healthconcerns include being a hazard to the applica-tor and farm workers, as well as to the consumerfrom high pesticide residues on the fruit. Typi-cally, those who spray the crop are not aware ofthe hazards of pesticides and do not use anypersonal protective equipment (Fig. 2). Theseproblems have been well documented in Ban-gladesh and other countries (bteggplant.cornell.edu/content/facts; bic.searca.org; Del Prado-Lu2015). In addition to environmental and healthconcerns, populations of EFSB have also devel-oped resistance to many foliar insecticides(Shirale et al. 2017).

EGGPLANT IN BANGLADESH

Bangladesh is a country of 147,570 km2 with apopulation of more than 165 million (Fig. 3).Vegetables are an important component of theBangladeshi diet and more than 90 vegetablesare cultivated. In terms of area and overall pro-duction, eggplant (brinjal) is the second-most-important vegetable grown by an estimated150,000 resource-poor farmers on 51,000 hathroughout Bangladesh. Brinjal constitutesabout 14.9% of winter vegetable production

Figure 1. Eggplant fruit and shoot borer damage innon-Bt brinjal. (Photograph from the personal collec-tion of Md. Arif Hossain.)

Figure 2. Bangladeshi farmer spraying an insecticidein non-Bt brinjal crop. (Photograph from the person-al collection of Md. Arif Hossain.)

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and 9.7% of summer production (BangladeshBureau of Statistics 2018).

DEVELOPMENT OF Bt EGGPLANT

As an alternative to intensive use of insecticides,the India-basedMaharashtraHybrid Seed Com-pany (Mahyco) inserted the cry1Ac gene, under

the control of the constitutive 35S CaMV pro-moter, into eggplant (termed “event” EE-1) tocontrol feeding damage by EFSB (Mahyco hashad a long-term relationship with Monsanto inIndia for producing Bt cotton. Monsanto wasrecently acquired by Bayer). The cry1A gene iswidely used in Bt cotton and has a long historyof safe use (ILSI CERA, 2010). Bt eggplant dem-

Figure 3. Map of Bangladesh. (Figure reprinted courtesy of childrenrights24@blogspot.com.)

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onstrated control of EFSB in contained green-house trials in India. In late 2003, a partnershipwas formed between Mahyco, Cornell Universi-ty, the United States Agency for InternationalDevelopment (USAID), and public sector part-ners in India, Bangladesh, and the Philippinesunder the Agricultural Biotechnology SupportProject II (ABSPII) (Shelton et al. 2017).Mahycodonated the EE-1 event to the Bangladesh Agri-cultural Research Institute (BARI), and Mahycoincorporated it into BARI-developed local open-pollinated (OP) varieties (OP lines allow farmersto save seed; however, BARI discourages seedsaving because of potential out-crossing, whichwould reduce the purity of the seed). BARI con-ducted confined field trials that demonstratedthe Bt varieties provided excellent control ofEFSB compared to non-Bt isolines (same varietybut without the Bt gene) (Fig. 4). Results fromtrials conducted in Bangladesh were used to de-velop the regulatory dossier that was submittedto Bangladeshi authorities for cultivation (seebelow).

The ABSPII project ended in 2014. A new3-year cooperative agreement was awarded in2015 under the USAID-funded Feed the FutureSouth Asia Eggplant Improvement Partnership(bteggplant.cornell.edu). The main objectiveswere to scale up the improved Bt eggplant toBangladeshi farmers, provide capacity buildingwithin the Bangladesh agricultural institutions,and work with the University of the Philippines-Los Baños to develop, submit, and shepherd a

regulatory dossier for the Philippines that meetsinternational standards. This article only focuseson Bangladesh.

THE INDIAN SITUATION

Much of the early ABSPII work on Bt eggplantwas centered in India, a country that producesnearly 25% of the world’s eggplants, grownmostly by smallholder farmers on a total of 1.3million acres but with relatively low productionbecause of infestation by EFSB (Krishna andQaim 2007). Beginning in 2004, Bt hybridswere tested in several states of India and per-formed well, leading Krishna and Qaim (2007)to write “several Bt hybrids have been tested inthe field and are likely to be commercialized inthe near future.”Unlike OP lines, hybrids do notbreed true the following year so new seed mustbe obtained yearly. However, farmers generallyprefer hybrids because of their enhanced yieldand quality.

After extensive field trials and safety evalua-tions by Indian regulatory bodies, Bt eggplantwas ready to be commercialized in India.However, antibiotech organizations, primarilyGreenpeace, had a different plan and for yearsfought against Bt eggplant in courts of law andin the court of public opinion. Greenpeace, theworld’s largest “environmental” organization,uses its considerable resources and influence toadvocate for a zero-tolerance for GM crops(Davidson 2008; Shelton 2015) (www.activistfacts

BA

Figure 4.Comparison of Uttara brinjal variety showing injury by (A) the eggplant fruit and shoot borer to non-Btbrinjal, and (B) lack of injury in Bt brinjal. (Photograph provided by M.J. Hossain, Feed the Future South AsiaEggplant Improvement Partnership.)

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.com/organizations/131-greenpeace).Greenpeacespent millions to prevent Bt eggplant from com-ing to market in India, with a large amount usedin legal costs (Shelton 2010). As a result, largelyof antibiotech activists’ efforts, the Indian Min-ister of the Environment and Forests, the last“gatekeeper” before Bt eggplant would be com-mercialized, decided to impose amoratoriumonBt eggplant on February 9, 2010, which remainstoday (Shelton 2010). India’s neighbor, Bangla-desh, which also produces eggplant, decided tomove forward.

Bt EGGPLANT BECOMES A REALITYIN BANGLADESH

After successfully breeding EE-1 into nine localOP varieties, BARI applied to the NationalTechnical Committee on Crop Biotechnology(NTCCB) for their release. Following the recom-mendation from theNTCCB, the application forrelease was forwarded to the National Commit-tee on BioSafety. The Bangladesh governmentgranted approval for release of four lines on Oc-tober 30, 2013 (Fig. 5). On January 22, 2014, Bteggplant seedlings were distributed to 20 farm-ers in four districts, making Bangladesh the firstdeveloping country to allow the commercial cul-tivation of a GE vegetable crop (Mondal 2018).

Bt brinjal was made available to growers fordemonstration trials. In 2014–2015, BARI pro-vided seeds or transplants to its On-FarmResearch Division) to conduct research/demon-stration trials on 108 farmer fields in 19 districts.In 2015–2016 and 2016–2017, demonstration

trials were conducted in 250 farmer fields in 25districts and 512 farmer fields in 36 districts,respectively. In 2017–2018, BARIprovided seedsto 569 farmers in 40 districts. In addition to dis-tribution by BARI, seeds were distributed tofarmers through theDepartment of AgriculturalExtension (DAE) to 6000 and 7001 farmers in2016–2017 and 2017–2018, respectively, and forsale through the Bangladesh Agricultural Devel-opment Corporation (BADC) to an additional17,950 farmers in 2018, making a total of 27,012farmers in 2018 (Mondal 2018; Shelton et al.2018). Fruit from these fields was sold in themarket and readily purchased by consumers. Be-cause the four lines are OP varieties, farmers cankeep their seed (or give or sell it to others) so theactual number of farmers growing Bt brinjal islikely much higher than the statistics reportedabove.

The initial application for Bt brinjal includ-ed nine varieties that would accommodate re-gional preferences, but only four were approvedin 2013. There are continuing efforts to releaseother varieties, especially ones that are resistantto bacterial wilt, a severe disease in Bangladesh.

CONTROLOF EFSB AND EFFECTSON NONTARGET ARTHROPODSIN BANGLADESH

A 2-year experiment (2016–2017) conducted byBARI scientists compared the four Bt varieties totheir isolines (same variety but without the Btgene) with and without insecticide treatments(Prodhan et al. 2018). Results indicated that

BARI Bt begun-1 BARI Bt begun-2 BARI Bt begun-3 BARI Bt begun-4

Figure 5. Four Bt brinjal lines currently commercialized in Bangladesh. (Photograph provided by M.J. Hossain,Feed the Future South Asia Eggplant Improvement Partnership.)

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the four Bt varieties had increased fruit produc-tion and minimal EFSB fruit infestation com-pared with their respective non-Bt isolines. Fruitinfestation for Bt varieties varied from 0% to2.27% in 2016, 0% in 2017, and was not signifi-cantly affected by the spray regime in either year.In contrast, fruit infestation in non-Bt linesreached 36.70% in 2016 and 45.51% in 2017,even with weekly spraying. It should also benoted that when a Bt fruit was classified as beinginfested, only a superficial blemish was seen andno live larvae were present.

Statistically similar densities of nontarget ar-thropods, including beneficial arthropods, weregenerally observed in both Bt and non-Bt vari-eties. An additional trial that focused on a singleBt variety and its isoline provided similar resultson infestation levels, with and without sprays,and showed no significant effects on nontargetarthropods. These results are similar to studiesconducted in the Philippines that demonstratedexcellent control of EFSB (Hautea et al. 2016)and lack of effect on nontarget organisms (Na-vasero et al. 2016).

ECONOMICS AND INSECTICIDE PATTERNSWHEN USING Bt BRINJAL

In an ex ante (before the event) study on adop-tion of Bt eggplant in Bangladesh (Islam andNorton 2007), it was estimated that adoptionof Bt eggplant would reduce insecticide use by80% and increase the gross profit margin bynearly 45%. Because Bt eggplant has been com-mercialized, these predictions can be comparedto actual results. In a study conducted by BARI,scientists in 35 districts during the 2016–2017cropping season using 505 Bt brinjal farmersand 350 non-Bt brinjal farmers, net returnsper hectare were $2151/ha for Bt brinjal as com-pared to $357/ha for non-Bt brinjal, a sixfolddifference (Rashid et al. 2018). This study alsoindicated that farmers saved 61% of the pesticidecost compared to non-Bt brinjal farmers andexperienced no losses because of EFSB. It is like-ly that pesticide reduction could be reducedfurther once farmers have more experience con-trolling EFSB with Bt eggplant, and thus eco-nomic returns would be even higher.

In the 2-year study conducted by Prodhanet al. (2018), an economic analysis revealed thatall Bt lines had higher gross returns than theirnon-Bt isolines. The nonsprayed, non-Bt iso-lines resulted in negative returns in most cases.Maximum fruit yield was obtained from sprayedplots compared to nonsprayed plots, indicatingthat other insects such as whiteflies, thrips, andmites (these arthropods are not affected byCry1Ac) can reduce plant vigor and subsequentfruit weight. Efforts are now underway to de-velop treatment guidelines for the complex of“sucking insects” that can reduce plant vigor.

SUSTAINING THE TECHNOLOGY

The long-term success of Bt brinjal in Bangla-desh depends on many biological and adminis-trative factors. Farmers depend on high qualityseed so a major effort in the project has been tobuild capacitywithinBARI to ensure that qualityseed—genetic purity, high viability, and expres-sion of Cry1Ac—is produced in adequateamounts tomeet growerdemand.Toaccomplishthis, the project has undertaken capacity build-ing efforts to train BARI scientists following themodel of “Excellence Through Stewardship,” alife-cycle approach to GE product management(www.excellencethroughstewardship.org).

Likewise, farmers are being trained on theunique aspects of Bt brinjal, mainly the require-ments to plant a refuge of non-Bt brinjal as bor-der rows and the need to manage other “suckinginsects.” A refuge is required as a strategy todelay the evolution of resistance by EFSB tothe Cry1Ac protein (Bates et al. 2005). Researchhas demonstrated that utilizing a refuge is vitalfor ensuring the durability of Bt technology(Tang et al. 2001). Therefore,monitoring farmeradoption of and compliance with refuge-plant-ing strategies is essential. Monitoring compli-ance of farmers who save or distribute theirsaved seed is even more challenging. Recentreports have highlighted the difficulty in moni-toring the planting of refuges in developingcountries (India and Brazil) compared to Aus-tralia and theUnited States (Carrière et al. 2019).An alternative to planting separate refuges is tomix Bt seeds with a proportion of non-Bt seeds

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in the same bag sold to growers, a strategy called“refuge in the bag” (RIB).While the RIB strategyessentially forces farmers to plant a refuge, it ismost suitable for insects that have limited move-ment between plants in the larval stage (Tanget al. 2001). Additional research is needed todetermine whether the RIB strategy will be suit-able for EFSB.

There is general agreement (Bates et al.2005) and experimental evidence (Zhao et al.2003) that plants expressingmultiple Bt proteinswill bemore durable for control of an insect pest.Thus, efforts should be focused on introducingeggplant varieties with multiple Bt genes, assoon as possible. Mahyco has developed a two-gene Bt event and there are discussions under-way about having access to it. Meanwhile, stud-ies have been completed to assess the baselinesusceptibility of populations of EFSB to theCry1Ac expressed in Bt eggplants, so anychanges in susceptibility over time can be deter-mined, and appropriate measures taken. Itshould also be noted that the four Bt varietiescurrently used are not suitable for the summerseason, a time when populations of EFSB are attheir highest. Fortuitously, this Bt crop-free pe-riod acts as another component of a resistancemanagement strategy. Other management strat-egies that are compatiblewith Bt plants are beingconsidered, including removing infested plants,using pheromone disruption, enhancing biolog-ical control, and using more selective insecti-cides. However, farmers are typically less willingto adopt such practices because of their in-creased labor and costs, compared to using Btplants (Shelton 2007).

COMMUNICATING ABOUT Bt BRINJAL

Although the BARI-USAID-Cornell-Sathguruproject focuses on Bt brinjal, it serves a largerrole because it is the first GE food crop to bereleased and widely adopted by farmers andconsumers in a developing country. Thus, ithas been in the spotlight globally for the last5 years. This project includes communicationefforts that focus on two major activities: (1)improved information sharing internally andexternally to promote science-based public

awareness of GE crops; and (2) support commu-nications, outreach, and knowledge sharing byfarmers, extension personnel, community or-ganizations, and communication practitioners.An important communication avenue isthe project’s website (Bteggplant.cornell.edu),which serves as the face of the project. This siteis actively maintained and contains print andaudiovisual materials for information sharingand awareness building. The project has devel-oped a strong social media presence on twitter(@Bt_eggplant) and through various blogs. Cur-rently, more than 4500 people follow posts andupdates through the Cornell Alliance for Sci-ence. Factual news from national and interna-tional articles are shared and comments andfeedback are monitored and responded towhen suitable. The website also contains severalvideos, including two produced by BARI, high-lighting the success of Bt brinjal in the field andendorsement by local farmers.

The project also works closely with the Cor-nell Alliance for Science (allianceforscience.cornell.edu), which provides factual informa-tion about agricultural biotechnology. The Alli-ance has enhanced capacity in social media thatbenefits the project in the short and long term.TheAlliance is leading efforts to expandbiotech-nology communication activities in Bangladeshto include other GE crops, for example, late-blight-resistant potatoes and nutritionally en-hanced rice. A new comprehensive communica-tion initiative fundedby theBill&MelindaGatesFoundation—Farming Future Bangladesh—hasbeen formed to create an empowered communi-ty of advocates to develop a sustainable enablingenvironment for biotechnology in Bangladesh.

WHY IS THE BANGLADESH Bt EGGPLANTPROJECT SUCCEEDING?

ABSPII was essential to the success of the Ban-gladesh Bt eggplant project because it identifieda strong need for the project, was well-supportedfinancially and programmatically by USAID,utilized effective partners in India (Sathguru),Bangladesh (BARI), and the United States (Cor-nell), and had the good fortune to have eventEE-1 generously donated by Mahyco. When

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ABSPII ended, the project continued with sup-port from USAID’s Feed the Future programstarted in 2015. In addition to these enablingfactors, it is essential to acknowledge that theproject has received political support from keyBangladeshi people since its inception. In con-trast to India where the project stopped becauseof political pressure from well-funded groupsopposed to biotechnology, Bangladesh receivedstrong support from the Bangladeshi Govern-ment, especially Prime Minister Sheikh Hasinaand the Honorable Agriculture Minister BegumMatia Chowdhury, MP. Chowdhury’s wordsfrom a Bt brinjal workshop held in March2017 in Bangladesh made the government’s po-sition clear:

Development of brinjal fruit and shoot insectresistant-Bt brinjal is a success story of localand foreign collaboration. We will be guided bythe science-based information, not by the non-scientific whispering of a section of people. Goodscience will move on its own course keeping theanti-science people down. As human beings, it isour moral obligation that all people in our coun-try should get food and not go to bed on an emptystomach. Biotechnology can play an importantrole in this effort.

In January 2019, Begum Matia Chowdhurywas replaced as the Agriculture Minister by Dr.M. Abdul Razzaque, awell-respected agricultur-

al scientist. At the time this article was written,efforts are underway to brief Dr. Razzaque inmore detail on the project and advocate for hissupport.

SECURING THE FUTURE OF Bt BRINJALIN BANGLADESH

The benefits of Bt brinjal are clear: increasedyield and economic return to farmers, decreasedrisk to humans and the environment, and sus-tainable production. Farmers are pleased withthe performance of Bt brinjal (Fig. 6). The ex-periences by farmers who have used Bt brinjalhave led to an increased demand for the seed,not only for the four current Bt lines but also foradditional Bt lines that would be suitable to oth-er regions or markets in Bangladesh. The ques-tion is whether the demand can be met by thecurrent system in which BARI is the soleproducer of Bt brinjal breeder seed, and BADCincreases the seed and sells it through their tra-ditional channels. So far, the private sector hasnot been allowed to produce Bt brinjal for dis-tribution in Bangladesh, although the linesgrown originated from the private sector’s (Ma-hyco) donated event, EE-1. The government ac-knowledges that the private sector should be aplayer in the future but has been hesitant toallow them to produce Bt brinjal for the time

Figure 6.Bangladeshi Bt brinjal farmer, ShahajahanAli, with harvested Bt brinjal. (Photograph from the personalcollection of Md. Arif Hossain.)

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being.Meanwhile, there are ongoing discussionsabout allowing the private sector to complementthe existing efforts of the government by pro-ducing their own hybrids.

In all but a few instances (e.g., virus-resistantpapaya), it has been the private sector that spentthe effort and considerable resources needed todevelop a transformed line (“event”), spentmillions on the regulatory process to have it ap-proved, developed varieties suitable for the loca-tion, distributed seeds to farmers, and providedstewardship practices (e.g., technology and com-munication) to enhance the product’s future.Bangladesh has had considerable assistancefrom the outside (Mahyco, Cornell, USAID) toget to the point where more than 27,000 ruralfarmers are producing Bt brinjal and selling it inthe local markets or to brokers who distribute tourban consumers.

Continued funding from the outside is un-certain so the Bangladesh government will haveto take more responsibility for the future of Btbrinjal. Our project has focused on capacitybuilding to help Bangladesh take on the varioustasks needed for successful seed production,distribution, cultivation, adoption, and refuge-monitoring of Bt brinjal. At the same time, weare promoting conversations between the Ban-gladeshi Government and the private sector tohelp coordinate the various activities needed.

There are certainly challenges ahead for thecontinued success of Bt brinjal in Bangladesh.However, these challenges can be met with adetailed strategic plan that ensures cooperationof Bangladeshi agencies and adequate resourcesfor Bt brinjal. Such planning will also reap ben-efits as other biotech crops are being developedin Bangladesh for the future. The private seedsector should play a critical role in such plans.

THE ROLE OF THE PRIVATE SECTORIN BANGLADESH

The seed sector in Bangladesh is changing.Thoughmost plant breeding in Bangladesh con-tinues to be in the public domain, private com-panies are becoming increasingly involved inplant breeding in response to the growing de-mand from farmers for good-quality and better-

performing varieties. Major, private-sector seedcompanies in Bangladesh include Supremeseeds, Lal Teer seeds, ACI seeds, GETCO, andothers, and NGO-operated seed enterprises, in-cluding the Bangladesh Rural AdvancementCommittee (BRAC), PROSHIKA, GrameenKrishi Foundation, Rangpur Dinajpur RuralService (RDRS), and others. These players areinvolved in trials of imported varieties of vege-tables and potatoes, development of improvedvarieties of indigenous vegetables through selec-tion and screening, development of hybrid veg-etables and maize, and seed production andmarketing. They also produce hybrid seeds ofrice and maize using imported parent lines.The private sector contributes in technology dis-semination and the education of farmers andtraders throughout the country. They have builtstrong marketing networks throughout thecountry. Private seed companies play majorroles in importing rice hybrids, seed potatoes,jute seeds, maize hybrids, and winter vegetableseeds where the public sector is generally inac-tive as a matter of policy.

Currently, in the private sector there aremore than 300 companies along with over22,000 registered seed dealers operating acrossBangladesh. They have a strong seed marketingnetwork throughout the country and strongcommitment to enhance their involvement inthis sector. Moreover, the growing engagementsof the private sector seed companies have result-ed in the engagement of thousands of contractfarmers in the formal seed production chain,leading to improved livelihoods for rural com-munities. USAID strongly believes in the impor-tance of the private sector becoming more in-volved in Bt brinjal seed production. The SeedWing of the Ministry of Agriculture and severaldonor-assisted projects have been acting as cat-alysts for building the capacity of the privatesector to play a greater role and take responsi-bility for achieving the seed policy goal of mak-ing quality seed available in timely ways to thefarmers of Bangladesh in cost-effective ways.The private sector has been provided with ade-quate representation in all bodies established bythe government for administering the seed in-dustry where the public/private/NGO sectors

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can discuss issues for the balanced and sustain-able growth of the seed sector.

With diminished resources available in thepublic sector over the last three decades, theprivate seed sector in Bangladesh has shownsignificant development and expansion. Achiev-ing an ideal partitioning of seed sector rolesamong private and public sectors under Bangla-desh agro-socio-economic conditions is noteasy given that the buying capacity of the farm-ers is limited and the government’s food securitygoals entail availability of quality seed at an af-fordable price. However, finding a balance willensure that safe and effective GE technologieswill reach resource-poor farmers in Bangladesh.

REPLICATING THE BANGLADESHI PROJECT

The results seen in Bangladesh demonstrate thatBt eggplant will result in economic benefits forfarmers and dramatically reduce health risks tohumans and the environment. Can this successbe replicated in other Asian countries whereEFSB ravages eggplants, especially in fieldsgrown by resource-poor farmers? Certainly,GE technology is available but it was availablein India before it was commercialized in Bangla-desh. History has shown that these two neigh-boring countries approached Bt eggplant differ-ently, with Bangladeshi leaders displaying thecourage to move forward based on the clearneed to help farmers reduce applications ofchemical sprays. In contrast, in spite of the sim-ilar needs of farmers in India and the recom-mendations of Indian scientists and regulatorsthat Bt brinjal is safe and effective, the IndianMinister of the Environment and Forests im-posed a moratorium on commercialization ofBt eggplant that remains today (Shelton 2010).It appears that the Indian minister based hisdecision not on science or farmers’ need butinstead on political pressure from groups againstGE crops. The consequence is that farmers inIndia continue to rely on chemical-based insec-ticide management practices to reduce damageby EFSB. Yet such a strategy is often ineffective,whereas Bt brinjal provides excellent controlwith minimal need for sprays for other pests(Prodhan et al. 2018).

In India, the intense use of pesticides onvegetables and other crops causes severe hazardsto farmers, consumers, and the environment(Grewal et al. 2017), but alternatives are avail-able. Bt cotton became available in India in 2002and remains the only GM product available toIndian farmers. Use of Bt cotton over a 10-yearperiod resulted in farmers reducing their pesti-cide use by at least 50% with the largest reduc-tions of 70% occurring with themost toxic typesof chemicals (Kouser and Qaim 2011). Thishelped avoid at least 2.4 million cases of pesti-cide poisoning, saving at least US$14 million inannual benefit costs. Could the same types ofbenefits occur in India if it were allowed to begrown in India, as it presently is in Bangladesh?

Time will tell whether Bt eggplant and otheruseful products of biotechnology will becomeavailable to benefit farmers, consumers, andthe environment in other countries, or willfear and misinformation impede their develop-ment and use? For the time being, Bt eggplant inBangladesh serves as a celebrated example of thesuccesses that can be achieved.

ACKNOWLEDGMENTS

The success of this project is the result of manyorganizations and people, including the Maha-rashtra Hybrid Seed Company (Mahyco) thatcreated the Bt eggplant “event” and incorporat-ed it into varieties used in Bangladesh, and whocontinue to provide important advice for theproject; the vision and support of the Govern-ment of Bangladesh; the Bangladesh Agricultur-al Research Institute (BARI) and its associatedagencies, including the Department of Agricul-tural Extension (DAE) and the Bangladesh Ag-ricultural Development Corporation (BADC);funding and advice provided by the UnitedStates Agency for International Development(USAID); personnel in Cornell’s College of Ag-riculture and Life Sciences International Pro-grams (CALS-IP) who managed the Agricultur-al Biotechnology Support Project II (ABSPII)and now manage the Feed the Future SouthAsia Eggplant Improvement Partnership; Sath-guru Management Consultants Pvt. Ltd.; andthe Cornell Alliance for Science. Without these

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partners, the project would not have been able tohave the impact it has had.

REFERENCES

Ali MI, Ali MS, Rahman MS. 1980. Field evaluation of wiltdisease and shoot and fruit borer attack of different cul-tivars of brinjal. Bangladesh J Agric Sci 7: 193–194.

Bangladesh Bureau of Statistics. 2018. Yearbook of Agricul-tural Statistics-2017, Bangladesh Bureau of Statistics(BBS) Statistics and Informatics Division (SID) Ministryof Planning Government of the People’s Republic of Ban-gladesh. www.bbs.gov.bd

Bates SL, Zhao JZ, Roush RT, Shelton AM. 2005. Insectresistance management in GM crops: Past, present andfuture. Nat Biotechnol 23: 57–62. doi:10.1038/nbt1056

BrookesG, Barfoot P. 2017.GMcrops: Global socio-economicand environmental impacts 1996–2015. PG EconomicsLtd., Dorchester, UK.

Carrière Y, Brown ZS, Downes SJ, Gujar G, Epstein G,Omoto C, Storer NP, Mota-Sanchez D, Jorgensen PS,Carroll SP. 2019. Governing evolution: A socioecologicalcomparison of resistance management for insecticidaltransgenic Bt crops among countries. Ambio doi:10.1007/s13280-019-01167-0

Davidson S. 2008. Forbidden fruit: Transgenic papaya inThailand. Plant Physiol 147: 487–493. doi:10.1104/pp.108.116913

Del Prado-Lu J. 2015. Insecticide residues in soil, water, andeggplant fruits and farmers’ health effects due to exposureto pesticides. Environ Health Prev Med 20: 53–62. doi:10.1007/s12199-014-0425-3

Fischhoff DA, Bowdishi KS, Perlak FJ, Marrone PG,McCor-mick SM, Niedermeyer JG, Dean DA, Kusano-KretzmerK, Mayer EJ, Rochester DE, et al. 1987. Insect toleranttransgenic tomato plants. Nat Biotechnol 5: 807–813.doi:10.1038/nbt0887-807

Grewal AS, Singlal A, Kamboj P, Dua JS. 2017. Pesticideresidues in food grains, vegetables and fruits: A hazardto human health. J Med Chem Toxicol 2: 40–46. doi:10.15436/2575-808x.17.1355

Hautea DM, Taylor LD, Masanga APL, Sison MLJ, NarcisoJO, Quilloy RB, Hautea RA, Shotkoski FA, Shelton AM.2016. Field performance of Bt eggplants (Solanummelon-gena L.) in the Philippines: Cry1Ac expression and con-trol of the eggplant fruit and shoot borer (Leucinodesorbonalis Guenée). PLoS ONE 11: e0157498. doi:10.1371/journal.pone.0157498

Heckel D. 2012. Learning the ABCs of Bt: ABC transportersand insect resistance toBacillus thuringiensisprovide cluesto a crucial step in toxin mode of action. Pestic BiochemPhysiol 104: 103–110. doi:10.1016/j.pestbp.2012.05.007

ILSI CERA. 2010. A review of the environmental safety of theCry1Ac protein. ILSI CERA, Washington, DC.

Islam SMF, Norton GW. 2007. Bt eggplant for fruit andshoot borer resistance in Bangladesh. In Economic andenvironmental benefits and costs of transgenic crops: Ex-ante assessment (ed. Ramasamy C, Selvaraj KN, NortonGW, Vijayaraghavan K), pp. 91–106. Tamil Nadu Agri-cultural University, Coimbatore, India.

James C. 2000. Global status of commercialized transgeniccrops: 2000. ISAAA Briefs, No. 21. ISAAA, Ithaca, NY.

James C. 2018. Global status of commercialized biotech/GMcrops: 2017. ISAAA Briefs, No. 53. ISAAA, Ithaca, NY.

Kennedy GG. 2008. Integration of insect-resistant genetical-ly modified crops within IPM programs. In Integration ofinsect-resistant genetically modified crops within IPM pro-grams (ed. Romeis J, Shelton A, Kennedy G), pp. 1–26.Springer Science+Business Media, New York.

Kouser S, Qaim M. 2011. Impact of Bt cotton on pesticidepoisoning in smallholder agriculture: A panel data anal-ysis. Ecol Econ 70: 2105–2113. doi:10.1016/j.ecolecon.2011.06.008

Krishna VV, Qaim M. 2007. Potential socieoeconomicimpacts of Bt eggplant in India. In Economic and envi-ronmental benefits and costs of transgenic crops: Ex-anteassessment (ed. Ramasamy C, et al.), pp. 57–71. TamilNadu Agricultural University, Coimbatore, India.

Mondal MRI. 2018. Success on Bt brinjal in Bangladesh.Asia-Pacific Consortium on Agricultural Biotechnologyand Bioresources Asia-Pacific Association of AgriculturalResearch Institutions. Bangkok, Thailand.

Naranjo SE, Ellsworth PC. 2009. Fifty years of the integratedcontrol concept: Moving the model and implementationforward inArizona.PestManag Sci 65: 1267–1286. doi:10.1002/ps.1861

Navasero MV, Candano RN, Hautea D, Hautea RA, Shot-koski FA, Shelton AM. 2016. Assessing potential impactof Bt eggplants on non-target arthropods in the Philip-pines. PLoS ONE 11: e0165190. doi:10.1371/journal.pone.165190

Prodhan MZH, Hasan MT, Chowdhury MMI, Alam MS,RahmanML, Azad AK, HossainMJ, Naranjo SE, SheltonAM. 2018. Bt Eggplant (Solanum melongena L.) in Ban-gladesh: Fruit production and control of eggplant fruitand shoot borer (Leucinodes orbonalis Guenee), effectson non-target arthropods and economic returns. PLoSONE 13: e0205713. doi:10.1371/journal.pone.0205713

Rashid MA, Hasan MK, Matin MA. 2018. Socio-economicperformance of Bt eggplant cultivation in Bangladesh.Bangladesh J Agric Res 43: 187–203. doi:10.3329/bjar.v43i2.37313

Romeis J, Shelton AM, Kennedy GG. 2008. Integration ofinsect-resistant genetically modified crops within IPM pro-grams, 441 pp. Springer, Dordrecht, Netherlands.

Romeis J, Naranjo SE, Meissle M, Shelton AM. 2019. Genet-ically engineered crops help support conservation biolog-ical control. Biol Control 130: 136–154. doi:10.1016/j.biocontrol.2018.10.001

Shelton AM. 2007. Considerations on the use of transgeniccrops for insect control. J Dev Stud 43: 890–900. doi:10.1080/00220380701384562

Shelton AM. 2010. The long road to commercialization of Btbrinjal (eggplant) in India. Crop Prot 29: 412–414. doi:10.1016/j.cropro.2010.02.016

Shelton AM. 2012. Genetically engineered vegetables ex-pressing proteins from Bacillus thuringiensis for insectresistance: Successes, disappointments, challenges andways to move forward. GM Crops Food 3: 175–183.doi:10.4161/gmcr.19762

A.M. Shelton et al.

12 Advanced Online Article. Cite this article as Cold Spring Harb Perspect Biol doi: 10.1101/cshperspect.a034678

Shelton AM. 2015. Tragic papaya: 12-part series.anthonymshelton.com/tragicpapaya

Shelton AM, Zhao JZ, Roush RT. 2002. Economic, ecologi-cal, food safety and social consequences of the deploy-ment of Bt transgenic plants. Annu Rev Entomol 47: 845–881. doi:10.1146/annurev.ento.47.091201.145309

Shelton AM, Olmstead DL, Burkness EC, Hutchison WD,Dively G, Welty C, Sparks AN. 2013. Multi-state trials ofBt sweet corn varieties for control of the corn earworm(Lepidoptera: Noctuidae). J Econ Entomol 106: 2151–2159. doi:10.1603/EC13146

Shelton AM, Hokanson KE, Hautea DM, Hossain MJ, Hos-sainMA, Paranjape V, Hautea RA,McCandless L, SarwerSH. 2017. Bt eggplant: A genetically engineered “minor”crop comes of age in Bangladesh and the Philippines. ISBNews Report, August.

Shelton AM, Hossain MJ, Paranjape V, Azad AK, RahamML, Khan ASMMR, Prodhan MZH, Rashid MA, Ma-jumder R, Hossain MA, et al. 2018. Bt eggplant projectin Bangladesh: History, present status, and future direc-tion. Front Bioeng Biotechnol 6: 106. doi:10.3389/fbioe.2018.00106

Shirale D, Patil M, Parimi S. 2017. Insecticide resistance infield populations of Leucinodes orbonalis (Lepidoptera:Crambidae) in India. Can Entomol 149: 399–407.doi:10.4039/tce.2017.3

Smith CM, Clement SL. 2012. Molecular bases of plantresistance to arthropods. Annu Rev Entomol 57: 309–328.

Stern V, Smith R, Van den Bosch R, Hagen K. 1959. Theintegration of chemical and biological control of the spot-ted alfalfa aphid: The integrated control concept. Hilgar-dia 29: 81–101. doi:10.3733/hilg.v29n02p081

Tang JD, Collins HL, Metz TD, Earle ED, Zhao J, Roush RT,Shelton AM. 2001. Greenhouse tests on resistance man-agement of Bt transgenic plants using refuge strategies. JEcon Entomol 94: 240–247. doi:10.1603/0022-0493-94.1.240

Zhao J, Cao J, Li Y, Collins HL, Roush RT, Earle ED, SheltonAM. 2003. Plants expressing two Bacillus thuringiensistoxins delay insect resistance compared to single toxinsused sequentially or in amosaic.Nat Biotechnol 21: 1493–1497. doi:10.1038/nbt907

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