Intended and unintended consequences of genetically modified crops – myth, fact and/or manageable outcomes?

Figures & data

Table 1. Number of commercial GM events by trait and crop species – summarised from ISAAA (2022).

Crop species	GM trait
	Abiotic stress tolerance	Altered growth/ yield	Disease resistance	Herbicide tolerance	Insect resistance	Modified product quality	Nematode tolerance	Pollination control system
Apple						3
Argentine canola				37		10		24
Bean			1
Chicory				3				3
Cotton				45	50	1
Cowpea					1
Creeping bentgrass				1
Eggplant					1
Eucalyptus		1
Flax				1
Lucerne				4		2
Maize	7	2		215	210	14		6
Melon						2
Papaya			4
Pineapple						1
Plum			1
Polish canola				4
Poplar					30
Potato			19	4	8	18
Rice				3	3	2
Safflower						2
Soybean	2	1		35	6	12	1
Squash			2
Sugar beet				3
Sugarcane	3
Sweet pepper			1
Tobacco				1		1
Tomato			1		1	9
Wheat	1			1
TOTAL	13	4	29	357	310	77	1	33

Table 2. Commercialised GM traits and their benefits – (also refer to Halford 2004; Nalluri and Karri 2020; Brookes and Barfoot 2020c for more detail).

Trait	Crop	% Total GM crops*	Intended consequence; efficacy and benefits	Reference
Glyphosate tolerance	Range of crops – maize, cotton, canola	–	Reduced use of a broad range of selective herbicides; replaced with used of broad spectrum herbicides such as glyphosate	Brookes and Barfoot (2020c)
	Canola	3.8	Weed control	Monjardino et al. (2005)
	Cotton	2.3	Improved weed control; reduced tillage	Ammann (2005); Dill et al. (2008)
	Maize	20.5	Weed control	Dill et al. (2008)
	Potato (Solanum tuberosum)	<0.1	Weed control	Bakhsh et al. (2020)
	Soybean	35.5	Improved weed control; reduced tillage	Ammann (2005); Dill et al. (2008)
	Sugar beet (Beta vulgaris)	0.2	Economic; weed control	Demont and Dillen (2008)
	Wheat	<0.1	Wheat transgenic event 33391 gave good glyphosate tolerance and yield	Zhou et al. (2003)
Insect pest resistance due to Bt Cry proteins	Range of crops – maize, cotton	–	Significantly reduced insecticide use	Betz et al. (2000); Brookes and Barfoot (2020c)
			30 pests controlled by Bt crops in Latin America	Blanco et al. (2016)
			Bt crops suppress pests, reduce pesticide use and increase farmer profits	Tabashnik and Carriere (2015)
	Egg plant, Brinjal (Solanum melongena)	<0.1	Control of major pests	Muhammad et al. (2020)
	Cotton	8.7	Control of cotton bollworm, Helicoverpa armigera	Lu et al. (2012); Jadhav et al. (2020)
	Cowpea (Vigna unguiculata)	<0.1	Control of bean pod borer, Maruca vitrata, which can cause 20–80% yield loss	Addae et al. (2020)
	Maize	19.3	Control of the European maize borer	Hutchison et al. (2010); Chvatalova (2020)
			Control of maize rootworms (Diabrotica spp.) – meta analysis of 10 publications	Pellegrino et al. (2018)
	Potatoes for industrial use and animal feed	<0.1	Control of Colorado potato beetle	James (2011); Kadoić Balaško et al. (2020)
	Rice	<0.1	Control of yellow stem borer (Scirpophaga incertulas), rice striped stem borer (Chilo suppressalis), pink stem borer (Sesamia inferens), and rice leaf roller (Cnaphalocrocis medinalis)	Bashir et al. (2004); Ramesh et al. (2004); Tang et al. (2006); Li et al. (2016)
	Soybean	9.8	Control of key soybean lepidopteran pests	Grossi-de-Sa et al. (2011); Bacalhau et al. (2020)
Combined herbicide tolerance, male sterility induction and insect resistance	Maize	<0.1	MON 87427 × MON 89034 × NK603 developed by conventional breeding	Venkatesh et al. (2014)
Disease resistance	Papaya (Carica papaya)	<0.1	Resistance to papaya ringspot virus	Lius et al. (1997); Gonsalves (1998); Gonsalves et al. (2007)
	Rice	<0.1	Resistance to bacterial blight (Xanthomonas oryzae)	Rani et al. (2008)
	Squash (Cucurbita sp.)	<0.1	Virus resistance	Fuchs and Gonsalves (1995); Tricoll et al. (1995)
Reduced lignin	Alfalfa (lucerne) (Medicago sativa)	<0.1	Improved forage digestibility	ISAAA (2017); Barros et al. (2019)
Drought tolerance	Maize	<0.1	Increase grain production in high-water-deficit environments	Chang et al. (2014)
			Use of event MON 87460 to improve drought tolerance though improved water use efficiency	ISAAA (2016)
Phytase expression	Maize	<0.1	Replace the additional P supplementation in monogastric animals, reducing the cost of feeding	Wang and Guan (2020)
Improved nutritional quality	Rice	<0.1	Expression of Vitamin A (Golden Rice); many years after development has become commercial in Philippines	Paine et al. (2005); Tang et al. (2009); Kim et al. (2010); Stone and Glover (2017); Sumangil (2022)

*Percentage of GM crop plantings in 2018 – from Figure 11 of Brookes and Barfoot (2020c); Those categorised as <0.1% were not included in the Brookes and Barfoot (2020c) review – so this level is below the lowest quoted by them.

Table 3. Calculated average percentage differences between GM and non-GM crops for important output variables from a meta-analysis by Klümper and Qaim (2014).

Output variable	Type of GM crop
	All GM crops	Insect resistant	Herbicide tolerant
Yield	+21**	+25**	+9*
Pesticide quantity used	−37**	−42**	+2
Pesticide cost	−39**	−43**	−25**
Total cost of production	+3	+5*	−7
Farmer profit	+68**	+69**	+64

*p < 0.05; **p < 0.01.

Table 4. Impact of changes in the use of herbicides and insecticides and subsequently calculated change in Environmental Impact Quotient (EIQ) from growing GM crops globally 1996–2015 (Slightly modified from Brookes and Barfoot 2017).

GM trait	Absolute change		Percentage change		Area planted (2015) (million ha)
	Weight active ingredient (million kg)	Field EIQ (million EIQ/ha units)	Active ingredient use	Environmental impact
Herbicide-tolerant soybean	+15	−8112	+0.5	−14	76.7
Herbicide and insect-resistant soybean	−3	−348	−4.3	−4	22.3
Herbicide-tolerant maize	−226	−7315	−8.4	−12	45.3
Herbicide-tolerant canola	−25	−854	−18.1	−30	8.6
Herbicide-tolerant cotton	−25	−629	−7.6	−10	4.2
Insect-resistant maize	−87	−3891	−53.3	−58	46.1
Insect-resistant cotton	−268	−11,949	−29.1	−31	21.5

Table 5. Summary of toxicology and health testing trials involving comparisons of GM crops and their non-GM equivalent indicating (A) some concerns and (B) little concern. Also refer to Magana-Gomez and Calderon de la Barca (2009) and DeFrancesco (2013) for additional information.

GM crop	Code or name and transgene	Type of study	Result compared to non-GM equivalent	Reference
A. Studies indicating some concern from toxicology and health testing trials
Maize	MON863 – Bt expressing Cry3Bb1	90 d rat feeding trial	Slight weight decrease and hepatorenal toxicity	Séralini et al. (2007)
	MON810 – Bt expressing Cry1Ab	30 and 90 d mouse feeding trial	Induced alterations in intestinal and peripheral immune response of weaning and old mice	Finamore et al. (2008)
	Bt	3 generations of feeding to rats	Some minimal histopathological changes in liver and kidney; no difference in organ weights	Kilic and Akay (2008)
	MON810 and MON863 – Bt expressing Cry1Ab and Cry3Bb1, respectively	Up to 90 d rat feeding trial	Negative impact on the function on kidneys and liver	de Vendomois et al. (2009)
	NK603 – glyphosate and AMPA tolerant	Up to 90 d rat feeding trial	Negative impact on the function on kidneys and liver	de Vendomois et al. (2009)
	NK603 – glyphosate tolerant	2-year feeding trial with rats	Females developed mammary tumours before controls; Males had four times more large palpable tumours starting earlier than in the control group	Séralini et al. (2014)
Peas (Pisum sativum)	Expression of Bean α-Amylase Inhibitor	Mice	Elicited specific immunoreactivity of these proteins	Prescott et al. (2005)
Potato	Expressing CryI gene	Mice	Mild changes are reported in the structural configuration of the ileum of mice	Fares and El-Sayed (1998)
	Expressing the lectin Galanthus nivalis agglutinin (GNA)	Rats	GNA expression was associated with significantly greater mucosal thickness of the stomach when compared with non-GM potato diets	Ewen and Pusztai (1999)
Soybean	Insertion of the bacterial CP4 EPSPS (5-enolpyruvylshikimate-3-phosphate synthase) gene conferring a high level of tolerance to glyphosate	Mice	GM soybean intake can nuclear features; however mechanisms responsible for this remain unclear	Malatesta et al. (2002)
	Insertion of the bacterial CP4 EPSPS (5-enolpyruvylshikimate-3-phosphate synthase) gene conferring a high level of tolerance to glyphosate	Mice	GM soybean can influence the pancreatic metabolism in mouse. Unfortunately, the reasons for such modifications remain so far unclear.	Malatesta et al. (2003)
	Glyphosate tolerance	Mice	In testes of GM-fed mice the number of perichromatin granules is higher and the nuclear pore density lower; could be due to presence of residual glyphosate	Vecchio et al. (2004)
	Not specified	Allergenic skin tests on humans	Skin test results of 49 patients showed 13 positive results to wild soybeans and 8 positive results to GMO soybeans	Yum et al. (2005)
	Insertion of the bacterial CP4 EPSPS (5-enolpyruvylshikimate-3-phosphate synthase) gene conferring a high level of tolerance to glyphosate	Mice	GM soybean intake can influence some liver features during ageing	Malatesta et al. (2008)
	Type GM soybean product, which was from Kyrgyzstan, was not described but PCR revealed the presence of the 35S + FMV promoter	30 d feeding trial with rats	Visible changes in the spleen characterised by an increase in size, average weight, and changes in its structure and colour	Egamberganovich (2020)
B. Studies indicating little concern from toxicology and health testing trials
Maize	MON863 – Bt expressing Cry3Bb1	90 d rat feeding trial	No adverse effects on health, body weight gain, food consumption, clinical pathology parameters (haematology, blood chemistry, urinalysis), organ weights, gross and microscopic appearance of tissues	Hammond et al. (2006)
	Maize line 1507, containing event DAS-Ø15Ø7-1 (1507), expressing Cry1F	Sub-chronic rat feeding trial over 90 days	No toxicologically significant differences for nutritional performance variables, clinical and neurobehavioral signs, ophthalmology, clinical pathology (haematology, clinical chemistry, coagulation, and urinalysis), organ weights, and gross and microscopic pathology	MacKenzie et al. (2007)
	Bt176 expressing Cry1Ab	Sheep feeding trial over 3 years	No adverse effects on their health or performance and that no horizontal gene transfer to ruminal microorganisms or animal tissues was detected	Trabalza-Marinucci et al. (2008)
	Stacked GM crop NK603 x MON810	Mice	Multi-generation study showed no statistically significant differences concerning parental body mass; however time-related negative reproductive effects of the GM maize; some effect on some metabolic pathways	Velimirov et al. (2008)
Mix of maize and soybean	GM maize – a combination of NK603, MON863 and MON810 genes (expressing the CP4 EPSPS, Cry 3Bb1 and Cry 1Ab proteins respectively), GM soybean was 100% Roundup Ready (expressing the CP4 EPSPS protein),	Long-term toxicology study of 22.7 weeks with isowean pigs	No differences between pigs fed the GM and non-GM diets for feed intake, weight gain, mortality and routine blood biochemistry measurements, but there was a slight increase in severe stomach inflammation amongst male and female pigs	Carman et al. (2013)
Potato	Cry1Ab gene	90 d rat feeding trial	No effect on growth rate, food intake, food efficiency, and general health status; no effect on serum concentrations of immunoglobulin A (IgA), IgG, IgM, orinterleukin 6	Rahnama et al. (2017)
Rice	Cry1Ab/Ac gene	50 and 110 d feed trial with rats	Exogenous gene and protein of the transgenic rice did not remain in the intestines	Xu et al. (2015b)
Soybean	Glyphosate tolerance	105 d feeding trials with mice and rats	No effect on growth, feeding value and histopathology of immune-related organs	Teshima et al. (2000)
	Glyphosate tolerance	90 d feeding trial with mice	No effect on macromolecular synthesis or cell growth and differentiation of testicular cell populations; no differences in litter sizes and body weights	Brake and Evenson (2004)
	Glyphosate tolerance	90 d feeding trial with rats	No differences for gross necropsy findings, haematological or urinalysis values and clinical serum parameters	Zhu et al. (2004)
	Not specified but did include 35S promoter and used soybean meal (solvent extracted)	Rabbits	35S CMV was not found in all samples of serum, heart, skeletal muscle, liver and kidney; no differences detected for body weight, fresh organ weights	Tudisco et al. (2006)
	Type of GM soybean not specified in abstract (article in Russian)	Comparison of third-generation female rats where both they and their mothers	No definite negative or positive impact of GM soy components on their physiological state compared with animals fed non-GM soybeans	Dolaychuk et al. (2013)
	MON87708 – dicamba tolerant	90-day sub-chronic toxicology study with rats	No indication of adverse effects associated with the consumption of dicamba-tolerant soybean	Wang et al. (2016)

Table 6. Summary of commercially released crops with transgenic nutritionally enhanced traits. Modified from Garg et al. (2018) and refer to McGloughlin (2010) and Jones (2015).

Crop	Nutritional trait	Company	Example reference
Canola	Phytate degradation (increase in available P)	BASF	Jones (2015)
	High lauric acid	Monsanto
Cassava (Manihot esculenta)	Iron	Biocassava Plus	Sayre et al. (2011)
	Beta-carotene
	Protein
Maize	Phytate degradation	Origin Agritech, China	Origin Agritech (2010)
	Lysine	Monsanto; Renessen	Frizzi et al. (2008); Tang et al. (2013)
Potato	Reduced amylose and increased amylopectin in starch granules	BASF	Karlsson et al. (2007)
Soybean	Oleic acid	Dupont; Monsanto	Zhang et al. (2014)
	γ-Linolenic Acid + stearidonic acid	Monsanto	Sato et al. (2004)

Table 7. Number of references indicating an impact or not of transgenic events on soil microorganisms due to pleiotropic and undetermined (pleiotropic or direct) effects. Taken from review by Turrini et al. (2015).

Transgenic trait or gene	Effects of transgenes expressed in different plant species on soil microorganisms
	Consistent impact	Transient impact	No effect
Bt genes for insect resistance	19	13	35
Resistance to pathogens	6	7	8
Tolerance to herbicides	6	5	8
Food nutritional benefits	9	3	6
Total	40	28	57

Table 8. The impacts of Bt proteins administered in the laboratory or measured in the field on soil invertebrates. Summarised from Icoz and Stotzky (2008).

Organism group	Number of studies	Study location*	Effect of cry proteins on organism
			Consistent impact	No effect
Earthworms	6	0.17	0	6
Woodlouse	4	0	1 (fed less)	3
Collembola	9	0.33	1 (changed feeding preference)	8
Nematodes	12	0.25	7	5

*Proportion of total studies undertaken that were in the field.

Table 9. Representative examples of potential benefits of genetic modification to crop plants – unexploited opportunities and future options – (also refer to Newell-McGloughlin 2008, McGloughlin 2010; Nalluri and Karri 2020 for further detail).

Trait	Crop	Intended consequence; efficacy and benefits	Reference
Herbicide tolerance	Rice	Engineering herbicide-resistant rice plants through CRISPR/Cas9-mediated homologous recombination of acetolactate synthase	Sun et al. (2016)
Insect pest resistance due to Bt Cry proteins	Broccoli (Brassica oleracea var. italica)	Pyramided Cry1Ac and Cry1C Bt genes control diamondback moth (Plutella xylostella) even those resistant to Cry1A or Cry1C toxins	Cao et al. (2002)
	Eggplant, Brinjal	Eggplant engineered to produce the Cry1Ac protein provided control of eggplant fruit and shoot borer (Leucinodes orbonalis) reducing need for pesticides	Hautea et al. (2016)
	Pigeon pea (Cajanus cajan)	Marker free transgenic events with Cry1Ac protein were found to confer resistant activity against pod borer Helicoverpa armigera larvae	Ganguly et al. (2022)
	Rice	Confers resistance to stem borers (Scirpophaga incertulas and Chilo suppressalis) and leaf folders; no direct negative effects on predators, parasitoids, non-lepidopterous herbivores, or soil invertebrates and soil health. But not commercialised by 2016.	Cohen et al. (2008); Fraiture et al. (2016)
	Table potatoes	Bt table potatoes were produced from 1996 to 2001 for control of Colorado potato beetle (Leptinotarsa decemlineata), due to complications connected with planting GM potatoes, new insecticide compounds, and rejection of the public	Grafius and Douches (2008); Kadoić Balaško et al. (2020)
Insect pest resistance	Cocoa (Theobroma cacao)	CRISPR/Cas9 used to insert the Non-Expressor of Pathogenesis-Related 3 (TcNPR3) gene, a suppressor of the defence response; resulted in increased resistance to infection with the cacao pathogen Phytophthora tropicalis	Fister et al. (2018)
	Rice	Plants expressing the cecropin A gene from the giant silk moth Hyalophora cecropia show enhanced resistance to Magnaporthe grisea, the causal agent of the rice blast disease	Coca et al. (2006)
	Various crops	Alternative GM technologies using lectins, protease inhibitors; multiple resistance genes; tissue specific expression	Christou et al. (2006)
Drought tolerance	Maize	ARGOS8 gene (a negative regulator of ethylene responses) variants of maize increased grain yield by five bushels per acre under flowering stress conditions and had no yield loss under well-watered conditions.	Shi et al. (2017)
	Multiple species	Expression of related cold shock proteins (CSPs) from bacteria, CspA from Escherichia coli and CspB from Bacillus subtilis, promotes stress adaptation	Castiglioni et al. (2008)
		Pathways and genes that can be manipulated through biotechnology to improve drought tolerance.	Deikman et al. (2012)
	Rice	GM rice containing the CaMsrB2 gene are superior in performance to non-GM rice in drought stress conditions	Racheal et al. (2020)
		Overexpression of stress responsive gene SNAC1 (STRESS-RESPONSIVE NAC 1) significantly enhances drought tolerance in transgenic rice	Hu et al. (2006)
		DREB 2A gene isolated from drought tolerant rice variety	Raj et al. (2015)
		Introduction of ZAT12 gene improved drought tolerance in rice	Devi et al. (2016)
	Wheat	NAC gene, TaNAC5D-2 a positive regulator of drought tolerance when silence in wheat increased water loss and decreased growth compared to control and negative control plants under drought stress	Ma et al. (2022)
Salinity tolerance	Chickpea (Cicer arietinum)	OsRuvB gene used to improve salinity tolerance	Preeti (2022)
	Goundnut (Arachis hypogaea)	Overexpression of AtNHX1 gene not only improved salt tolerance but also drought tolerance in transgenic groundnut	Asif et al. (2011)
	Mulberry (Morus indica)	Improved salinity tolerance due to the transgene barley hva1 gene which encodes a group 3 LEA protein and is induced by ABA and water deficit conditions	Lal et al. (2008)
	Rice	Overexpression of stress responsive gene SNAC1 (STRESS-RESPONSIVE NAC 1) significantly enhances salinity tolerance in transgenic rice	Hu et al. (2006)
		Over expression of glyoxalase II that increased further upon salt stress, reflecting the upregulation of endogenous glyoxalase II.	Singla-Pareek et al. (2008)
		At the seedling stage, the salinity tolerance of T2 homozygous mutant lines with the OsRR22 gene was significantly enhanced compared to wild-type plants	Zhang et al. (2019)
	Tobacco (Nicotiana tabacum)	Three salt-stress induced hemp NAC genes, especially CsNAC1 and CsNAC3, played significant roles in the improvement of salt tolerance in transgenic tobacco	Liu et al. (2022)
	Tomato	Improved tolerance to salinity	Zhang and Blumwald (2011)
Nutritional and health benefits (also refer Garg et al. 2018)	Banana (Musa spp.)	6-fold increase in iron accumulation and a 4.5-fold increase in the zinc levels were showed in transgenic banana expressing ferritin constitutively	Chen et al. (2014)
		The fifth exon of the lycopene epsilon-cyclase (LCYϵ) gene was targeted to develop the β-carotene-enriched Cavendish banana cultivar	Kaur et al. (2020)
		Increased level of provitamin A	Namanya et al. (2020)
	Barley	Cisgenic lines with a single extra copy of the HvPAPhy resulted in 2.6- and 2.8-fold increases in phytase activities, that should result in increasing bioavailable phosphate from the stored phytic acid	Holme et al. (2012)
	Brassica	Modification of a fatty acid desaturase 2 gene (FAD2) leading to an increase in the content of oleic acid compared with that in wild-type seeds.	Okuzaki et al. (2018)
		Expression of omega-3 oils – eicosapentaenoic acid (EPA) and docosahexanoic acid (DHA) – in canola	Napier et al. (2019b)
	Camelina sativa	Expression of omega-3 oils – eicosapentaenoic acid (EPA) and docosahexanoic acid (DHA)	Usher et al. (2015)
	Cassava	Modification of starch quality in storage roots	Bull et al. (2018)
	Cereals	Express additional proteins which are rich in lysine and/or methionine	Shewry (2007); Galili and Amir (2013)
	Maize	Use of zinc-finger to alter the inositol phosphate profile in developing seeds	Shukla et al. (2009)
	Peanut	Positive AhFAD2 mutant lines displayed a 0.5–2 fold increase in the oleic acid content compared with non-transgenic controls	Wen et al. (2018)
	Potato	Silencing two asparagine synthetase genes through ‘all-native DNA’ Transformation of potato. French fries and potato chips accumulated as little as 5% of the acrylamide (toxic and carcinogenic – Perera et al. 2021) present in wild-type controls.	Rommens et al. (2008)
	Rice	High expression of amylose (resistant starch) rice through CRISPR/Cas9-mediated editing of SBEIIb, to reduce risk of non-infectious diseases	Sun et al. (2017)
		Knock out of Inositol 1,3,4-trisphosphate 5/6-kinase6 (IPTK6) gene results in low phytic acid expression; also impairs plant growth and reproduction. Chemically induced mutants were created that reduce phytic acid but did not affect plant growth	Kim and Tai (2014); Jiang et al. (2019)
		Carotenoid biosynthesis cassette at two genomic sites produces high carotenoid content in the seeds and no detectable penalty in morphology or yield.	Dong et al. (2020)
	Soybean	Reduced levels of polyunsaturated fats by introducing mutations in two fatty acid desaturase 2 genes (FAD2-1A and FAD2-1B), which in the seed convert the monounsaturated fat, oleic acid, to the polyunsaturated fat, linoleic acid	Haun et al. (2014)
	Tomato	Gene edited tomatoes produce more vitamin D	Li et al. (2022)
		Incorporation of bacterial carotenoid gene encoding the enzyme phytoene desaturase increased β-carotene content by threefold, up to 45% of the total carotenoid content.	Römer et al. (2000)
		A ‘purple’ tomato, highly enriched with anthocyanins, was produced by the ectopic expression of two selected transcription factors from the ornamental flower snapdragon. These fruits also prolonged the life of cancer-susceptible mice.	Gonzali et al. (2009)
		Expression of a yeast S-adenosylmethionine decarboxylase gene (ySAMdc; Spe2) fused with a ripening-inducible E8 promoter to enhance phytonutrient content, juice quality, and vine life	Mehta et al. (2002)
	Wheat	Manipulation the expression of high molecular weight glutenin subunits – associated with elasticity of dough for bread making and cause of celiac in humans	Blechl and Anderson (1996); Alvarez et al. (2000)
		Two sgRNAs were used to target a conserved region adjacent to the coding sequence a-gliadin genes, leading to a strong reduction in a-gliadins – low gluten wheat	Sanchez-Leon et al. (2018)
Disease resistance	Various crops	Although technically useful solutions now exist for providing transgenic disease resistance, very few new GM crops with these traits have been introduced to the global market	Collinge et al. (2010); Collinge and Sarrocco (2022)
		Integrating antiviral genes of virus origin into plant genomes to aid resistance to viral pathogens	Niraula and Fondong (2021)
	Banana	Constitutive expression of apoptosis-inhibition-related genes in transgenic banana ‘Lady Finger’ showed high levels of resistance to Fusarium wilt	Chen et al. (2014)
		Increased resistance to banana bacterial wilt	Namanya et al. (2020)
		Improvement of B genome germplasm to produce hybrids of plantain (Musa balbisiana) leading to inactivation of endogenous banana streak virus (eBSV) in the B genome	Tripathi et al. (2019)
	Citrus (Citrus spp,)	Canker (Xanthomonas citri subsp. citri) resistant plants developed through editing the susceptibility gene promoter	Peng et al. (2017)
	Papaya	Control of Papaya ringspot virus disease	Azad et al. (2014)
	Peanut	Resistance to Phaeoisariopsis personata leaf spot disease	Vasavirama and Kirti (2012)
	Pearl millet (Pennisetum glaucum)	Expression of antifungal protein AFP from the mould Aspergillus giganteus provided protection against Puccinia substriata and Sclerospora graminicola	Girgi et al. (2006)
	Potato	Resistance to late blight, caused by the oomycete Phytophthora infestans which is only controlled by fungicides	Haverkort et al. (2009); Zhu et al. (2013)
	Rice	Constitutively express chitinase (chi11) and β−1,3-glucanase (GLU) genes improved sheath blight disease (Rhizoctonia solani) resistance	Kannan et al. (2017)
		Disruption of transcription activator-like effectors (TALEs) reduced expression of a family of putative sugar transporter genes conferring improved resistance to bacterial blight (Xanthomonas oryzae pv. oryzae)	Xu et al. (2019); Oliva et al. (2019)
		Editing at least one allele of the coilin gene considerably increased the resistance of the edited lines to infection with the potato virus Y and their tolerance to salt and osmotic stress.	Makhotenko et al. (2019)
	Tea (Camellia sinensis)	Resistance to blister blight disease (Exobasidium vexans)	Singh et al. (2018)
	Tomato	Trans-acting small interfering RNA control of geminiviruses – tomato leaf curl Gujarat virus	Singh et al. (2015)
	Wheat	Broad-spectrum resistance to powdery mildew (Blumeria graminis f. sp. tritici)	Wang et al. (2014); Zhang et al. (2017)
	Wheat and barley	synergistic efforts in traditional breeding and biotechnology should provide wheat and barley growers with Fusarium head blight resistant cultivars	Dahleen et al. (2001)
Processability	Banana	Downregulated ACC synthase or ACC oxidase can change banana fruit ripening process significantly through reduced ethylene production, decreased respiration rate, and delayed the change of peel colour in the ripening stage	Chen et al. (2014)
	Potato	RNA interference (RNAi) technology to silence the vacuolar invertase gene, which encodes a protein that breaks down sucrose to glucose and fructose; improves storage quality at low temperature	Clasen et al. (2016)
	Sugarcane (Saccharum officinarum)	Increased yields of directly fermentable glucose from lignocellulosic biomass for ethanol production	Jung et al. (2012)
Reproductive efficiency	Maize	I–CreI homing endonuclease from Chlamydomonas reinhardti to gene edit and provide male sterility for hybrid breeding systems	Djukanovic et al. (2013)
Phenotype	Banana	Used CRISPR/Cas9 to edit the MaGA20ox2 genes in ‘Gros Michel’ and obtained semi-dwarf mutants	Shao et al. (2020)
Lignin modification	Lucerne	Lignin in lucerne can be influenced using a transcription factor (Zhou et al. 2022).	Zhou et al. (2022)

Table 10. Interpretation of evidence for benefits accruing from intended outcomes and risks associated with unintended outcomes from use of GM crops.

Consequence		Degree of benefit	Degree of concern	Comment
Intended	Economic gains	High in most circumstances, including small-holder farmers	Concern often associated with perceived or real influence of multinationals	Despite some concerns in developing countries
	Improved productivity and crop yield performance	High in most circumstances, including small-holder farmers	Lower increases in non-stressed environments	If correct, then should contribute to lower deforestation
	Crop management	Improved ease of management		Farmer uptake of GM crops has not been impeded by the need for on-farm management changes
	Crop product composition and quality	Good – generally compositional equivalence between GM and non-GM crops	Inferior lint quality in Bt cotton in Burkina Faso	Has been a fundamental component of human health safety assessment; use of Bt GM maize has reduced mycotoxins
	Agronomic and phenotypic plant traits	Generally equivalent to non-GM, even for stacked GM traits		Even when agronomic differences are documented they are unlikely to have biological or ecological significance
	Improved resistance to pests	High – achieving opportunities not available to non-GM breeding	A wider pool of insect control protein is required	Bt proteins generally not harmful to humans and target specific insect pests
	Reduced use of synthetic chemical	Generally a reduction through and Bt GM crops can be applied as part of Integrated Pest Management	Potential for producing resistance in weeds and insect pests	Negative environmental impacts reduced through using herbicide tolerant and pest resistant GM crops
	Lower C footprint and reduced greenhouse gas emissions	Some reduction through reduced use of petrochemicals	No data for reduction in nitrous oxide emissions	More than likely a positive outcome
	Reduced tillage	While tillage may have been reduced the environmental impact is unclear	Assumptions for reduced C lost from soil need tested	Some claims do not seem to be valid in practice

Consequence		Consideration	Degree of concern	Comment
Unintended	Increased herbicide tolerance of weeds	Developing herbicide resistance of weeds is not just restricted to GM crops	Can occur	Mitigated by use of herbicides with different modes of action
	Overuse of herbicides to mitigate tolerance of weed to glyphosate	GM herbicide tolerance continues to deliver significant economic and environmental gains	Can occur	Herbicide use seems to have increased on both GM and non-GM crops
	Overuse of pesticides to mitigate developing pest resistance to Bt proteins	Important pests have also become resistant to synthetic insecticides requiring increased use	Can occur	Bt crops may/can resolve some pest resistance issues to synthetic chemistry
	Secondary pests becoming dominant	If this does occur, then requires an to pest management systems	Rare, but some secondary pests can increase in importance	Secondary pests can be controlled by going back to insecticides, use of biological pest control, or use of multiple stacked Bt genes
	Impacts on non-target organisms	Often shown in laboratory studies but rarely in the field; some misleading evidence has been used to suggest non-target effects of Bt proteins	Bt proteins unlikely to have negative impacts on nontarget organisms	Bt proteins target a specific pest whereas crop protection chemicals may affect both intended target and beneficial organisms
	Resistant insect populations	Important pests have also become resistant to synthetic	Can occur	Mitigated by used of refuges and potentially use of a wider range of insecticidal proteins
	Increased weediness	GM arable crops are unlikely to survive for long outside cultivation	Unlikely
	Impacts on biodiversity	Preservation of biodiversity in natural ecosystems is of paramount importance	In GM crops biodiversity effects are low compared to use of synthetic chemistry	Any negative or less than desirable effect requires improvements in use are required through better management
	Transgene escape to wild populations	Many GM crop plants have wild relatives with which they can hybridise	Will occur in outcrossing species	Need to consider whether the transfer of transgene would be detrimental and create an unmanageable risk
	Impacts on rhizosphere microbes	Impacts depend on transformation events, experimental conditions and taxa analysed	Some effects but not necessarily deleterious	Synthetic chemistry can also potentially disrupt N cycling
	Horizontal gene flow	Poses negligible risks to human health or the environment	Unlikely and remote, requires effective selection processes	Gene transfer between bacteria in the phytosphere may be part of their evolutionary development
	Increased antibiotic resistance	GM plants most often uses an antibiotic-resistant gene as a selectable marker	Unlikely and remote, requires effective selection processes	No documented evidence of this occurring in 25 years of GM crop use
	Gene transfer to consumers	Gene transfer from GM food is no different from gene transfer in non-GM feeds	Highly unlikely
	Unintended compounds produced	Concern for unintended toxins being produced by GM crops has been proposed	No evidence of this	Bt GM maize may reduce mycotoxins in stored grain as a health benefit

Consequence		Consideration	Degree of concern	Comment
Other considerations	Safety of GM feed for animals	Testing for feed safety carried out on actual animal species	Equivalent to non-GM when fed to a range of farmed animals	Majority of GM crops are used for animal feed
	Food safety	No consistent and reliably documented issues; regulatory testing might that food from GM crops may be safer than that from non-GM crops	No human ailments, sickness or deaths linked to GM crops; food safety issues debates result from laboratory tests using rodents	Has caused polarised views; many non-GM crops also contain potentially toxic or allergenic compounds; level of risks to human health from foods contaminated with pathogens is far greater than from GM crops
	Nutritive value	Crops could be genetically transformed to improve their nutritive quality and value in an attempt to eradicate or lessen the impact of malnutrition	Nutritive value the same as non-GM or higher when selected for beneficial nutritive compounds	Uptake of nutritionally enhanced GM crops has been relatively slow for both animal and human consumption

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