La ingeniería genética , también llamada modificación genética o manipulación genética , es la manipulación directa de los genes de un organismo mediante la biotecnología . Es un conjunto de tecnologías que se utilizan para cambiar la composición genética de las células, incluida la transferencia de genes dentro y a través de los límites de las especies para producir organismos mejorados o nuevos . El nuevo ADN se obtiene aislando y copiando el material genético de interés utilizando métodos de ADN recombinante o sintetizando artificialmente el ADN. Una construccióngeneralmente se crea y se utiliza para insertar este ADN en el organismo huésped. Paul Berg fabricó la primera molécula de ADN recombinante en 1972 combinando ADN del virus del mono SV40 con el virus lambda . Además de insertar genes , el proceso se puede utilizar para eliminar o " anular " genes. El nuevo ADN se puede insertar al azar, o dirigido a una parte específica del genoma . [1]
Un organismo que se genera mediante ingeniería genética se considera genéticamente modificado (GM) y la entidad resultante es un organismo genéticamente modificado (OGM). El primer OGM fue una bacteria generada por Herbert Boyer y Stanley Cohen en 1973. Rudolf Jaenisch creó el primer animal transgénico cuando insertó ADN extraño en un ratón en 1974. La primera compañía en enfocarse en ingeniería genética, Genentech, fue fundada en 1976 y inició la producción de proteínas humanas. La insulina humana genéticamente modificada se produjo en 1978 y las bacterias productoras de insulina se comercializaron en 1982. Los alimentos genéticamente modificados se han vendido desde 1994, con el lanzamiento del tomate Flavr Savr . El Flavr Savr fue diseñado para tener una vida útil más larga, pero la mayoría de los cultivos transgénicos actuales se modifican para aumentar la resistencia a los insectos y herbicidas. GloFish , el primer OMG diseñado como mascota, se vendió en Estados Unidos en diciembre de 2003. En 2016 se vendió salmón modificado con una hormona del crecimiento.
La ingeniería genética se ha aplicado en numerosos campos, como la investigación, la medicina, la biotecnología industrial y la agricultura. En la investigación, los OGM se utilizan para estudiar la función y expresión de genes a través de experimentos de pérdida de función, ganancia de función, rastreo y expresión. Al eliminar los genes responsables de ciertas condiciones, es posible crear organismos modelo animales de enfermedades humanas. Además de producir hormonas, vacunas y otros medicamentos, la ingeniería genética tiene el potencial de curar enfermedades genéticas a través de la terapia génica . Las mismas técnicas que se utilizan para producir medicamentos también pueden tener aplicaciones industriales, como la producción de enzimas para detergente para ropa, quesos y otros productos.
El auge de los cultivos comercializados genéticamente modificados ha proporcionado beneficios económicos a los agricultores en muchos países diferentes, pero también ha sido la fuente de la mayor parte de la controversia en torno a la tecnología. Esto ha estado presente desde su uso temprano; los primeros ensayos de campo fueron destruidos por activistas anti-transgénicos. Aunque existe un consenso científico de que los alimentos actualmente disponibles derivados de cultivos transgénicos no representan un riesgo mayor para la salud humana que los alimentos convencionales, la seguridad alimentaria transgénica es una de las principales preocupaciones de los críticos. El flujo de genes , el impacto en organismos no objetivo, el control del suministro de alimentos y los derechos de propiedad intelectual también se han planteado como problemas potenciales. Estas preocupaciones han llevado al desarrollo de un marco regulatorio, que comenzó en 1975. Ha llevado a un tratado internacional, el Protocolo de Cartagena sobre Bioseguridad , que fue adoptado en 2000. Los países individuales han desarrollado sus propios sistemas regulatorios con respecto a los OGM, con el las diferencias más marcadas se producen entre los EE. UU. y Europa.
Descripción general
La ingeniería genética es un proceso que altera la estructura genética de un organismo eliminando o introduciendo ADN . A diferencia de la cría tradicional de animales y plantas , que implica hacer múltiples cruces y luego seleccionar el organismo con el fenotipo deseado , la ingeniería genética toma el gen directamente de un organismo y lo entrega al otro. Esto es mucho más rápido, se puede usar para insertar cualquier gen de cualquier organismo (incluso los de diferentes dominios ) y evita que también se agreguen otros genes indeseables. [4]
La ingeniería genética podría reparar trastornos genéticos graves en los seres humanos reemplazando el gen defectuoso por uno que funciona. [5] Es una herramienta importante en la investigación que permite estudiar la función de genes específicos. [6] Se han obtenido medicamentos, vacunas y otros productos de organismos diseñados para producirlos. [7] Se han desarrollado cultivos que ayudan a la seguridad alimentaria aumentando el rendimiento, el valor nutricional y la tolerancia a las tensiones ambientales. [8]
El ADN se puede introducir directamente en el organismo huésped o en una célula que luego se fusiona o hibrida con el huésped. [9] Esto se basa en técnicas de ácido nucleico recombinante para formar nuevas combinaciones de material genético hereditario seguido de la incorporación de ese material ya sea indirectamente a través de un sistema de vectores o directamente a través de microinyección , macroinyección o microencapsulación . [10]
La ingeniería genética normalmente no incluye la reproducción tradicional, la fertilización in vitro , la inducción de poliploidía , la mutagénesis y las técnicas de fusión celular que no utilizan ácidos nucleicos recombinantes o un organismo modificado genéticamente en el proceso. [9] Sin embargo, algunas definiciones amplias de ingeniería genética incluyen la cría selectiva . [10] La clonación y la investigación con células madre , aunque no se consideran ingeniería genética, [11] están estrechamente relacionadas y la ingeniería genética se puede utilizar en ellas. [12] La biología sintética es una disciplina emergente que lleva la ingeniería genética un paso más allá al introducir material sintetizado artificialmente en un organismo. [13] En este nuevo campo se fabrica ADN sintético como el Sistema de información genética expandido artificialmente y el ADN de Hachimoji .
Las plantas, los animales o los microorganismos que se han modificado mediante la ingeniería genética se denominan organismos modificados genéticamente o OMG. [14] Si se agrega material genético de otra especie al hospedador, el organismo resultante se denomina transgénico . Si se utiliza material genético de la misma especie o una especie que puede reproducirse naturalmente con el huésped, el organismo resultante se denomina cisgénico . [15] Si se utiliza la ingeniería genética para eliminar material genético del organismo objetivo, el organismo resultante se denomina organismo knockout . [16] En Europa, la modificación genética es sinónimo de ingeniería genética, mientras que en los Estados Unidos de América y Canadá la modificación genética también se puede utilizar para referirse a métodos de reproducción más convencionales. [17] [18] [19]
Historia
Los seres humanos han alterado los genomas de las especies durante miles de años mediante la reproducción selectiva o la selección artificial [20] : 1 [21] : 1 en contraste con la selección natural . Más recientemente, la reproducción por mutaciones ha utilizado la exposición a sustancias químicas o radiación para producir una alta frecuencia de mutaciones aleatorias, con fines de reproducción selectiva. La ingeniería genética como manipulación directa del ADN por humanos fuera de la reproducción y las mutaciones solo existe desde la década de 1970. El término "ingeniería genética" fue acuñado por primera vez por Jack Williamson en su ciencia ficción novela Isla del Dragón , publicado en 1951 [22] - un año antes el papel del ADN en la herencia fue confirmada por Alfred Hershey y Martha Chase , [23] y dos años antes James Watson y Francis Crick demostraron que la molécula de ADN tiene una estructura de doble hélice, aunque el concepto general de manipulación genética directa se exploró de forma rudimentaria en la historia de ciencia ficción de Stanley G. Weinbaum de 1936, Proteus Island . [24] [25]
En 1972, Paul Berg creó las primeras moléculas de ADN recombinante combinando el ADN del virus del mono SV40 con el del virus lambda . [26] En 1973, Herbert Boyer y Stanley Cohen crearon el primer organismo transgénico insertando genes de resistencia a los antibióticos en el plásmido de una bacteria Escherichia coli . [27] [28] Un año después, Rudolf Jaenisch creó un ratón transgénico mediante la introducción de ADN extraño en su embrión, convirtiéndolo en el primer animal transgénico del mundo [29] Estos logros generaron preocupación en la comunidad científica sobre los riesgos potenciales de la ingeniería genética, que se discutieron por primera vez en profundidad en la Conferencia de Asilomar en 1975. Una de las principales recomendaciones de esta reunión fue que se debería establecer la supervisión gubernamental de la investigación del ADN recombinante hasta que la tecnología se considere segura. [30] [31]
En 1976 , Herbert Boyer y Robert Swanson fundaron Genentech , la primera empresa de ingeniería genética, y un año después la empresa produjo una proteína humana ( somatostatina ) en E. coli . Genentech anunció la producción de insulina humana modificada genéticamente en 1978. [32] En 1980, la Corte Suprema de los Estados Unidos en el caso Diamond v. Chakrabarty dictaminó que se podía patentar la vida genéticamente alterada. [33] La insulina producida por bacterias fue aprobada para su liberación por la Administración de Drogas y Alimentos (FDA) en 1982. [34]
En 1983, una empresa de biotecnología, Advanced Genetic Sciences (AGS) solicitó la autorización del gobierno de los EE. UU. Para realizar pruebas de campo con la cepa sin hielo de Pseudomonas syringae para proteger los cultivos de las heladas, pero los grupos ambientalistas y los manifestantes retrasaron las pruebas de campo durante cuatro años con impugnaciones legales. [35] En 1987, la cepa sin hielo de P. syringae se convirtió en el primer organismo genéticamente modificado (OMG) que se liberó al medio ambiente [36] cuando se roció con él un campo de fresas y un campo de patatas en California. [37] Ambos campos de prueba fueron atacados por grupos activistas la noche antes de que ocurrieran las pruebas: "El primer sitio de prueba del mundo atrajo al primer destructor de campo del mundo". [36]
Las primeras pruebas de campo de plantas modificadas genéticamente se realizaron en Francia y los Estados Unidos en 1986, las plantas de tabaco fueron diseñadas para ser resistentes a los herbicidas . [38] La República Popular de China fue el primer país en comercializar plantas transgénicas, introduciendo un tabaco resistente a virus en 1992. [39] En 1994 Calgene obtuvo la aprobación para comercializar el primer alimento modificado genéticamente , el Flavr Savr , un tomate modificado para tener una vida útil más larga. [40] En 1994, la Unión Europea aprobó el tabaco modificado para ser resistente al herbicida bromoxinil , convirtiéndolo en el primer cultivo modificado genéticamente comercializado en Europa. [41] En 1995, la Papa Bt fue aprobada como segura por la Agencia de Protección Ambiental , después de haber sido aprobada por la FDA, convirtiéndola en el primer cultivo productor de pesticidas aprobado en los EE. UU. [42] En 2009, se cultivaron comercialmente 11 cultivos transgénicos en 25 países, los mayores de los cuales por área cultivada fueron los Estados Unidos, Brasil, Argentina, India, Canadá, China, Paraguay y Sudáfrica. [43]
En 2010, los científicos del Instituto J. Craig Venter crearon el primer genoma sintético y lo insertaron en una célula bacteriana vacía. La bacteria resultante, llamada Mycoplasma laboratorium , podría replicarse y producir proteínas. [44] [45] Cuatro años más tarde, esto dio un paso más cuando se desarrolló una bacteria que replicaba un plásmido que contenía un par de bases único , creando el primer organismo diseñado para usar un alfabeto genético expandido. [46] [47] En 2012, Jennifer Doudna y Emmanuelle Charpentier colaboraron para desarrollar el sistema CRISPR / Cas9 , [48] [49] una técnica que puede usarse para alterar fácil y específicamente el genoma de casi cualquier organismo. [50]
Proceso
La creación de un OMG es un proceso de varios pasos. Los ingenieros genéticos primero deben elegir qué gen desean insertar en el organismo. Esto está impulsado por el objetivo del organismo resultante y se basa en investigaciones anteriores. Se pueden llevar a cabo exámenes genéticos para determinar genes potenciales y luego se pueden usar más pruebas para identificar a los mejores candidatos. El desarrollo de microarrays , transcriptómica y secuenciación del genoma ha facilitado mucho la búsqueda de genes adecuados. [51] La suerte también juega su papel; el gen listo para redondear se descubrió después de que los científicos notaron una bacteria que prosperaba en presencia del herbicida. [52]
Aislamiento y clonación de genes
El siguiente paso es aislar el gen candidato. Se abre la célula que contiene el gen y se purifica el ADN. [53] El gen se separa mediante el uso de enzimas de restricción para cortar el ADN en fragmentos [54] o la reacción en cadena de la polimerasa (PCR) para amplificar el segmento del gen. [55] Estos segmentos pueden luego extraerse mediante electroforesis en gel . Si el gen elegido o el genoma del organismo donante se ha estudiado bien, es posible que ya sea accesible desde una biblioteca genética . Si se conoce la secuencia de ADN , pero no hay copias disponibles del gen, también se puede sintetizar artificialmente . [56] Una vez aislado, el gen se liga a un plásmido que luego se inserta en una bacteria. El plásmido se replica cuando las bacterias se dividen, lo que garantiza la disponibilidad de copias ilimitadas del gen. [57]
Antes de que el gen se inserte en el organismo objetivo, debe combinarse con otros elementos genéticos. Estos incluyen una región promotora y terminadora , que inician y finalizan la transcripción . Se agrega un gen marcador seleccionable , que en la mayoría de los casos confiere resistencia a los antibióticos , por lo que los investigadores pueden determinar fácilmente qué células se han transformado con éxito. El gen también se puede modificar en esta etapa para una mejor expresión o efectividad. Estas manipulaciones se llevan a cabo mediante técnicas de ADN recombinante , como digestiones de restricción , ligaciones y clonación molecular. [58]
Inserción de ADN en el genoma del huésped
Hay una serie de técnicas que se utilizan para insertar material genético en el genoma del huésped. Algunas bacterias pueden absorber ADN extraño de forma natural . Esta capacidad se puede inducir en otras bacterias a través del estrés (por ejemplo, descarga térmica o eléctrica), lo que aumenta la permeabilidad de la membrana celular al ADN; El ADN recuperado puede integrarse con el genoma o existir como ADN extracromosómico . El ADN generalmente se inserta en células animales mediante microinyección , donde se puede inyectar a través de la envoltura nuclear de la célula directamente en el núcleo o mediante el uso de vectores virales . [59]
Los genomas de plantas se pueden diseñar mediante métodos físicos o mediante el uso de Agrobacterium para la entrega de secuencias alojadas en vectores binarios de T-DNA . En las plantas el ADN se inserta a menudo usando Agrobacterium mediada por transformación , [60] aprovechando la Agrobacterium s T-ADN de secuencia que permite la inserción natural del material genético en las células vegetales. [61] Otros métodos incluyen la biolística , donde las partículas de oro o tungsteno se recubren con ADN y luego se inyectan en las células de las plantas jóvenes, [62] y la electroporación , que implica el uso de una descarga eléctrica para hacer que la membrana celular sea permeable al ADN plasmídico.
Como solo una sola célula se transforma con material genético, el organismo debe regenerarse a partir de esa única célula. En las plantas, esto se logra mediante el uso de cultivo de tejidos . [63] [64] En los animales, es necesario asegurarse de que el ADN insertado esté presente en las células madre embrionarias . [65] Las bacterias consisten en una sola célula y se reproducen clonalmente, por lo que la regeneración no es necesaria. Los marcadores seleccionables se utilizan para diferenciar fácilmente las células transformadas de las no transformadas. Estos marcadores suelen estar presentes en el organismo transgénico, aunque se han desarrollado varias estrategias que pueden eliminar el marcador seleccionable de la planta transgénica madura. [66]
Se realizan más pruebas mediante PCR, hibridación Southern y secuenciación de ADN para confirmar que un organismo contiene el nuevo gen. [67] Estas pruebas también pueden confirmar la ubicación cromosómica y el número de copias del gen insertado. La presencia del gen no garantiza que se expresará a niveles apropiados en el tejido diana, por lo que también se utilizan métodos que buscan y miden los productos génicos (ARN y proteína). Estos incluyen hibridación Northern , RT-PCR cuantitativa , Western blot , inmunofluorescencia , ELISA y análisis fenotípico. [68]
El nuevo material genético puede insertarse aleatoriamente dentro del genoma del huésped o dirigirse a una ubicación específica. La técnica de selección de genes utiliza la recombinación homóloga para realizar los cambios deseados en un gen endógeno específico . Esto tiende a ocurrir con una frecuencia relativamente baja en plantas y animales y generalmente requiere el uso de marcadores seleccionables . La frecuencia de la selección de genes se puede mejorar enormemente mediante la edición del genoma . La edición del genoma utiliza nucleasas diseñadas artificialmente que crean rupturas específicas de doble hebra en lugares deseados del genoma y utilizan los mecanismos endógenos de la célula para reparar la ruptura inducida por los procesos naturales de recombinación homóloga y unión de extremos no homóloga . Hay cuatro familias de nucleasas diseñadas: meganucleasas , [69] [70] nucleasas con dedos de zinc , [71] [72] nucleasas efectoras de tipo activador de la transcripción (TALEN), [73] [74] y el sistema Cas9-guideRNA (adaptado de CRISPR ). [75] [76] TALEN y CRISPR son los dos más utilizados y cada uno tiene sus propias ventajas. [77] Los TALEN tienen una mayor especificidad de objetivo, mientras que CRISPR es más fácil de diseñar y más eficiente. [77] Además de mejorar el direccionamiento de genes, las nucleasas manipuladas pueden usarse para introducir mutaciones en genes endógenos que generan un gen knockout . [78] [79]
Aplicaciones
La ingeniería genética tiene aplicaciones en medicina, investigación, industria y agricultura y se puede utilizar en una amplia gama de plantas, animales y microorganismos. A las bacterias , los primeros organismos que se modifican genéticamente, se les puede insertar ADN plasmídico que contiene nuevos genes que codifican medicamentos o enzimas que procesan alimentos y otros sustratos . [80] [81] Las plantas han sido modificadas para protección contra insectos, resistencia a herbicidas, resistencia a virus, mejor nutrición, tolerancia a presiones ambientales y la producción de vacunas comestibles . [82] La mayoría de los OMG comercializados son plantas de cultivo resistentes a insectos o tolerantes a herbicidas. [83] Los animales genéticamente modificados se han utilizado para la investigación, animales modelo y la producción de productos agrícolas o farmacéuticos. Los animales genéticamente modificados incluyen animales con genes eliminados , mayor susceptibilidad a las enfermedades , hormonas para un crecimiento adicional y la capacidad de expresar proteínas en la leche. [84]
Medicamento
La ingeniería genética tiene muchas aplicaciones en la medicina que incluyen la fabricación de medicamentos, la creación de animales modelo que imitan las condiciones humanas y la terapia génica . Uno de los primeros usos de la ingeniería genética fue la producción masiva de insulina humana en bacterias. [32] Esta aplicación Ahora se ha aplicado a humanos hormonas de crecimiento , folículo estimulante hormonas (por infertilidad tratamiento), albúmina humana , anticuerpos monoclonales , factores antihemofílicos , vacunas y muchos otros medicamentos. [85] [86] Los hibridomas de ratón , células fusionadas para crear anticuerpos monoclonales , se han adaptado mediante ingeniería genética para crear anticuerpos monoclonales humanos. [87] En 2017, la FDA de EE. UU. Aprobó la ingeniería genética de receptores de antígenos quiméricos en las propias células T de un paciente como tratamiento para la leucemia linfoblástica aguda por cáncer . Se están desarrollando virus genéticamente modificados que aún pueden conferir inmunidad, pero carecen de las secuencias infecciosas . [88]
La ingeniería genética también se utiliza para crear modelos animales de enfermedades humanas. Los ratones genéticamente modificados son el modelo animal modificado genéticamente más común. [89] Se han utilizado para estudiar y modelar el cáncer (el oncomouse ), la obesidad, las enfermedades cardíacas, la diabetes, la artritis, el abuso de sustancias, la ansiedad, el envejecimiento y la enfermedad de Parkinson. [90] Las curas potenciales se pueden probar con estos modelos de ratón. También se han criado cerdos modificados genéticamente con el objetivo de aumentar el éxito del trasplante de órganos de cerdo a humano . [91]
La terapia génica es la ingeniería genética de los seres humanos , generalmente mediante la sustitución de genes defectuosos por otros eficaces. Se han realizado investigaciones clínicas que utilizan terapia génica somática con varias enfermedades, incluida la SCID ligada al cromosoma X , [92] la leucemia linfocítica crónica (LLC), [93] [94] y la enfermedad de Parkinson . [95] En 2012, Alipogene tiparvovec se convirtió en el primer tratamiento de terapia génica aprobado para uso clínico. [96] [97] En 2015, se usó un virus para insertar un gen sano en las células de la piel de un niño que padecía una enfermedad cutánea poco común, la epidermólisis ampollosa , con el fin de crecer, y luego injertar piel sana en el 80 por ciento de la piel del niño. cuerpo afectado por la enfermedad. [98]
La terapia génica de la línea germinal daría como resultado que cualquier cambio fuera heredable, lo que ha generado preocupaciones dentro de la comunidad científica. [99] [100] En 2015, CRISPR se utilizó para editar el ADN de embriones humanos no viables , [101] [102] los principales científicos de las principales academias mundiales para pedir una moratoria sobre las ediciones del genoma humano heredables. [103] También existe la preocupación de que la tecnología pueda usarse no solo para el tratamiento, sino para mejorar, modificar o alterar la apariencia, adaptabilidad, inteligencia, carácter o comportamiento de un ser humano. [104] La distinción entre curación y mejora también puede ser difícil de establecer. [105] En noviembre de 2018, He Jiankui anunció que había editado los genomas de dos embriones humanos para intentar desactivar el gen CCR5 , que codifica un receptor que el VIH usa para ingresar a las células. Dijo que las gemelas, Lulu y Nana, habían nacido unas semanas antes. Dijo que las niñas todavía llevaban copias funcionales de CCR5 junto con CCR5 discapacitado ( mosaicismo ) y aún eran vulnerables al VIH. El trabajo fue ampliamente condenado como poco ético, peligroso y prematuro. [106] Actualmente, la modificación de la línea germinal está prohibida en 40 países. Los científicos que realizan este tipo de investigación a menudo dejan que los embriones crezcan durante unos días sin permitir que se desarrollen hasta convertirse en un bebé. [107]
Researchers are altering the genome of pigs to induce the growth of human organs to be used in transplants. Scientists are creating "gene drives", changing the genomes of mosquitoes to make them immune to malaria, and then looking to spread the genetically altered mosquitoes throughout the mosquito population in the hopes of eliminating the disease.[108]
Research
Genetic engineering is an important tool for natural scientists, with the creation of transgenic organisms one of the most important tools for analysis of gene function.[109] Genes and other genetic information from a wide range of organisms can be inserted into bacteria for storage and modification, creating genetically modified bacteria in the process. Bacteria are cheap, easy to grow, clonal, multiply quickly, relatively easy to transform and can be stored at -80 °C almost indefinitely. Once a gene is isolated it can be stored inside the bacteria providing an unlimited supply for research.[110] Organisms are genetically engineered to discover the functions of certain genes. This could be the effect on the phenotype of the organism, where the gene is expressed or what other genes it interacts with. These experiments generally involve loss of function, gain of function, tracking and expression.
- Loss of function experiments, such as in a gene knockout experiment, in which an organism is engineered to lack the activity of one or more genes. In a simple knockout a copy of the desired gene has been altered to make it non-functional. Embryonic stem cells incorporate the altered gene, which replaces the already present functional copy. These stem cells are injected into blastocysts, which are implanted into surrogate mothers. This allows the experimenter to analyse the defects caused by this mutation and thereby determine the role of particular genes. It is used especially frequently in developmental biology.[111] When this is done by creating a library of genes with point mutations at every position in the area of interest, or even every position in the whole gene, this is called "scanning mutagenesis". The simplest method, and the first to be used, is "alanine scanning", where every position in turn is mutated to the unreactive amino acid alanine.[112]
- Gain of function experiments, the logical counterpart of knockouts. These are sometimes performed in conjunction with knockout experiments to more finely establish the function of the desired gene. The process is much the same as that in knockout engineering, except that the construct is designed to increase the function of the gene, usually by providing extra copies of the gene or inducing synthesis of the protein more frequently. Gain of function is used to tell whether or not a protein is sufficient for a function, but does not always mean it's required, especially when dealing with genetic or functional redundancy.[111]
- Tracking experiments, which seek to gain information about the localisation and interaction of the desired protein. One way to do this is to replace the wild-type gene with a 'fusion' gene, which is a juxtaposition of the wild-type gene with a reporting element such as green fluorescent protein (GFP) that will allow easy visualisation of the products of the genetic modification. While this is a useful technique, the manipulation can destroy the function of the gene, creating secondary effects and possibly calling into question the results of the experiment. More sophisticated techniques are now in development that can track protein products without mitigating their function, such as the addition of small sequences that will serve as binding motifs to monoclonal antibodies.[111]
- Expression studies aim to discover where and when specific proteins are produced. In these experiments, the DNA sequence before the DNA that codes for a protein, known as a gene's promoter, is reintroduced into an organism with the protein coding region replaced by a reporter gene such as GFP or an enzyme that catalyses the production of a dye. Thus the time and place where a particular protein is produced can be observed. Expression studies can be taken a step further by altering the promoter to find which pieces are crucial for the proper expression of the gene and are actually bound by transcription factor proteins; this process is known as promoter bashing.[113]
Industrial
Organisms can have their cells transformed with a gene coding for a useful protein, such as an enzyme, so that they will overexpress the desired protein. Mass quantities of the protein can then be manufactured by growing the transformed organism in bioreactor equipment using industrial fermentation, and then purifying the protein.[114] Some genes do not work well in bacteria, so yeast, insect cells or mammalians cells can also be used.[115] These techniques are used to produce medicines such as insulin, human growth hormone, and vaccines, supplements such as tryptophan, aid in the production of food (chymosin in cheese making) and fuels.[116] Other applications with genetically engineered bacteria could involve making them perform tasks outside their natural cycle, such as making biofuels,[117] cleaning up oil spills, carbon and other toxic waste[118] and detecting arsenic in drinking water.[119] Certain genetically modified microbes can also be used in biomining and bioremediation, due to their ability to extract heavy metals from their environment and incorporate them into compounds that are more easily recoverable.[120]
In materials science, a genetically modified virus has been used in a research laboratory as a scaffold for assembling a more environmentally friendly lithium-ion battery.[121][122] Bacteria have also been engineered to function as sensors by expressing a fluorescent protein under certain environmental conditions.[123]
Agriculture
One of the best-known and controversial applications of genetic engineering is the creation and use of genetically modified crops or genetically modified livestock to produce genetically modified food. Crops have been developed to increase production, increase tolerance to abiotic stresses, alter the composition of the food, or to produce novel products.[125]
The first crops to be released commercially on a large scale provided protection from insect pests or tolerance to herbicides. Fungal and virus resistant crops have also been developed or are in development.[126][127] This makes the insect and weed management of crops easier and can indirectly increase crop yield.[128][129] GM crops that directly improve yield by accelerating growth or making the plant more hardy (by improving salt, cold or drought tolerance) are also under development.[130] In 2016 Salmon have been genetically modified with growth hormones to reach normal adult size much faster.[131]
GMOs have been developed that modify the quality of produce by increasing the nutritional value or providing more industrially useful qualities or quantities.[130] The Amflora potato produces a more industrially useful blend of starches. Soybeans and canola have been genetically modified to produce more healthy oils.[132][133] The first commercialised GM food was a tomato that had delayed ripening, increasing its shelf life.[134]
Plants and animals have been engineered to produce materials they do not normally make. Pharming uses crops and animals as bioreactors to produce vaccines, drug intermediates, or the drugs themselves; the useful product is purified from the harvest and then used in the standard pharmaceutical production process.[135] Cows and goats have been engineered to express drugs and other proteins in their milk, and in 2009 the FDA approved a drug produced in goat milk.[136][137]
Other applications
Genetic engineering has potential applications in conservation and natural area management. Gene transfer through viral vectors has been proposed as a means of controlling invasive species as well as vaccinating threatened fauna from disease.[138] Transgenic trees have been suggested as a way to confer resistance to pathogens in wild populations.[139] With the increasing risks of maladaptation in organisms as a result of climate change and other perturbations, facilitated adaptation through gene tweaking could be one solution to reducing extinction risks.[140] Applications of genetic engineering in conservation are thus far mostly theoretical and have yet to be put into practice.
Genetic engineering is also being used to create microbial art.[141] Some bacteria have been genetically engineered to create black and white photographs.[142] Novelty items such as lavender-colored carnations,[143] blue roses,[144] and glowing fish[145][146] have also been produced through genetic engineering.
Regulación
The regulation of genetic engineering concerns the approaches taken by governments to assess and manage the risks associated with the development and release of GMOs. The development of a regulatory framework began in 1975, at Asilomar, California.[147] The Asilomar meeting recommended a set of voluntary guidelines regarding the use of recombinant technology.[30] As the technology improved the US established a committee at the Office of Science and Technology,[148] which assigned regulatory approval of GM food to the USDA, FDA and EPA.[149] The Cartagena Protocol on Biosafety, an international treaty that governs the transfer, handling, and use of GMOs,[150] was adopted on 29 January 2000.[151] One hundred and fifty-seven countries are members of the Protocol and many use it as a reference point for their own regulations.[152]
The legal and regulatory status of GM foods varies by country, with some nations banning or restricting them, and others permitting them with widely differing degrees of regulation.[153][154][155][156] Some countries allow the import of GM food with authorisation, but either do not allow its cultivation (Russia, Norway, Israel) or have provisions for cultivation even though no GM products are yet produced (Japan, South Korea). Most countries that do not allow GMO cultivation do permit research.[157] Some of the most marked differences occurring between the US and Europe. The US policy focuses on the product (not the process), only looks at verifiable scientific risks and uses the concept of substantial equivalence.[158] The European Union by contrast has possibly the most stringent GMO regulations in the world.[159] All GMOs, along with irradiated food, are considered "new food" and subject to extensive, case-by-case, science-based food evaluation by the European Food Safety Authority. The criteria for authorisation fall in four broad categories: "safety", "freedom of choice", "labelling", and "traceability".[160] The level of regulation in other countries that cultivate GMOs lie in between Europe and the United States.
Region | Regulators | Notes |
---|---|---|
US | USDA, FDA and EPA[149] | |
Europe | European Food Safety Authority[160] | |
Canada | Health Canada and the Canadian Food Inspection Agency[161][162] | Regulated products with novel features regardless of method of origin[163][164] |
Africa | Common Market for Eastern and Southern Africa[165] | Final decision lies with each individual country.[165] |
China | Office of Agricultural Genetic Engineering Biosafety Administration[166] | |
India | Institutional Biosafety Committee, Review Committee on Genetic Manipulation and Genetic Engineering Approval Committee[167] | |
Argentina | National Agricultural Biotechnology Advisory Committee (environmental impact), the National Service of Health and Agrifood Quality (food safety) and the National Agribusiness Direction (effect on trade)[168] | Final decision made by the Secretariat of Agriculture, Livestock, Fishery and Food.[168] |
Brazil | National Biosafety Technical Commission (environmental and food safety) and the Council of Ministers (commercial and economical issues)[168] | |
Australia | Office of the Gene Technology Regulator (oversees all GM products), Therapeutic Goods Administration (GM medicines) and Food Standards Australia New Zealand (GM food).[169][170] | The individual state governments can then assess the impact of release on markets and trade and apply further legislation to control approved genetically modified products.[170] |
One of the key issues concerning regulators is whether GM products should be labeled. The European Commission says that mandatory labeling and traceability are needed to allow for informed choice, avoid potential false advertising[171] and facilitate the withdrawal of products if adverse effects on health or the environment are discovered.[172] The American Medical Association[173] and the American Association for the Advancement of Science[174] say that absent scientific evidence of harm even voluntary labeling is misleading and will falsely alarm consumers. Labeling of GMO products in the marketplace is required in 64 countries.[175] Labeling can be mandatory up to a threshold GM content level (which varies between countries) or voluntary. In Canada and the US labeling of GM food is voluntary,[176] while in Europe all food (including processed food) or feed which contains greater than 0.9% of approved GMOs must be labelled.[159]
Controversia
Critics have objected to the use of genetic engineering on several grounds, including ethical, ecological and economic concerns. Many of these concerns involve GM crops and whether food produced from them is safe and what impact growing them will have on the environment. These controversies have led to litigation, international trade disputes, and protests, and to restrictive regulation of commercial products in some countries.[177]
Accusations that scientists are "playing God" and other religious issues have been ascribed to the technology from the beginning.[178] Other ethical issues raised include the patenting of life,[179] the use of intellectual property rights,[180] the level of labeling on products,[181][182] control of the food supply[183] and the objectivity of the regulatory process.[184] Although doubts have been raised,[185] economically most studies have found growing GM crops to be beneficial to farmers.[186][187][188]
Gene flow between GM crops and compatible plants, along with increased use of selective herbicides, can increase the risk of "superweeds" developing.[189] Other environmental concerns involve potential impacts on non-target organisms, including soil microbes,[190] and an increase in secondary and resistant insect pests.[191][192] Many of the environmental impacts regarding GM crops may take many years to be understood and are also evident in conventional agriculture practices.[190][193] With the commercialisation of genetically modified fish there are concerns over what the environmental consequences will be if they escape.[194]
There are three main concerns over the safety of genetically modified food: whether they may provoke an allergic reaction; whether the genes could transfer from the food into human cells; and whether the genes not approved for human consumption could outcross to other crops.[195] There is a scientific consensus[196][197][198][199] that currently available food derived from GM crops poses no greater risk to human health than conventional food,[200][201][202][203][204] but that each GM food needs to be tested on a case-by-case basis before introduction.[205][206][207] Nonetheless, members of the public are less likely than scientists to perceive GM foods as safe.[208][209][210][211]
En la cultura popular
Genetic engineering features in many science fiction stories.[212] Frank Herbert's novel The White Plague described the deliberate use of genetic engineering to create a pathogen which specifically killed women.[212] Another of Herbert's creations, the Dune series of novels, uses genetic engineering to create the powerful but despised Tleilaxu.[213] Films such as The Island and Blade Runner bring the engineered creature to confront the person who created it or the being it was cloned from. Few films have informed audiences about genetic engineering, with the exception of the 1978 The Boys from Brazil and the 1993 Jurassic Park, both of which made use of a lesson, a demonstration, and a clip of scientific film.[214][215] Genetic engineering methods are weakly represented in film; Michael Clark, writing for The Wellcome Trust, calls the portrayal of genetic engineering and biotechnology "seriously distorted"[215] in films such as The 6th Day. In Clark's view, the biotechnology is typically "given fantastic but visually arresting forms" while the science is either relegated to the background or fictionalised to suit a young audience.[215]
Ver también
- Biological engineering
- Modifications (genetics)
- RNA editing#Therapeutic mRNA Editing
- Mutagenesis (molecular biology technique)
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The labeling should include objective information to the effect that a food or feed consists of, contains or is produced from GMOs. Clear labeling, irrespective of the detectability of DNA or protein resulting from the genetic modification in the final product, meets the demands expressed in numerous surveys by a large majority of consumers, facilitates informed choice and precludes potential misleading of consumers as regards methods of manufacture or production.
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(3) Traceability requirements for GMOs should facilitate both the withdrawal of products where unforeseen adverse effects on human health, animal health or the environment, including ecosystems, are established, and the targeting of monitoring to examine potential effects on, in particular, the environment. Traceability should also facilitate the implementation of risk management measures in accordance with the precautionary principle. (4) Traceability requirements for food and feed produced from GMOs should be established to facilitate accurate labeling of such products.
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We have reviewed the scientific literature on GE crop safety for the last 10 years that catches the scientific consensus matured since GE plants became widely cultivated worldwide, and we can conclude that the scientific research conducted so far has not detected any significant hazard directly connected with the use of GM crops. The literature about Biodiversity and the GE food/feed consumption has sometimes resulted in animated debate regarding the suitability of the experimental designs, the choice of the statistical methods or the public accessibility of data. Such debate, even if positive and part of the natural process of review by the scientific community, has frequently been distorted by the media and often used politically and inappropriately in anti-GE crops campaigns.
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Currently available transgenic crops and foods derived from them have been judged safe to eat and the methods used to test their safety have been deemed appropriate. These conclusions represent the consensus of the scientific evidence surveyed by the ICSU (2003) and they are consistent with the views of the World Health Organization (WHO, 2002). These foods have been assessed for increased risks to human health by several national regulatory authorities (inter alia, Argentina, Brazil, Canada, China, the United Kingdom and the United States) using their national food safety procedures (ICSU). To date no verifiable untoward toxic or nutritionally deleterious effects resulting from the consumption of foods derived from genetically modified crops have been discovered anywhere in the world (GM Science Review Panel). Many millions of people have consumed foods derived from GM plants – mainly maize, soybean and oilseed rape – without any observed adverse effects (ICSU).
- ^ Ronald P (May 2011). "Plant genetics, sustainable agriculture and global food security". Genetics. 188 (1): 11–20. doi:10.1534/genetics.111.128553. PMC 3120150. PMID 21546547.
There is broad scientific consensus that genetically engineered crops currently on the market are safe to eat. After 14 years of cultivation and a cumulative total of 2 billion acres planted, no adverse health or environmental effects have resulted from commercialization of genetically engineered crops (Board on Agriculture and Natural Resources, Committee on Environmental Impacts Associated with Commercialization of Transgenic Plants, National Research Council and Division on Earth and Life Studies 2002). Both the U.S. National Research Council and the Joint Research Centre (the European Union's scientific and technical research laboratory and an integral part of the European Commission) have concluded that there is a comprehensive body of knowledge that adequately addresses the food safety issue of genetically engineered crops (Committee on Identifying and Assessing Unintended Effects of Genetically Engineered Foods on Human Health and National Research Council 2004; European Commission Joint Research Centre 2008). These and other recent reports conclude that the processes of genetic engineering and conventional breeding are no different in terms of unintended consequences to human health and the environment (European Commission Directorate-General for Research and Innovation 2010).
- ^ But see also: Domingo JL, Giné Bordonaba J (May 2011). "A literature review on the safety assessment of genetically modified plants". Environment International. 37 (4): 734–42. doi:10.1016/j.envint.2011.01.003. PMID 21296423.
In spite of this, the number of studies specifically focused on safety assessment of GM plants is still limited. However, it is important to remark that for the first time, a certain equilibrium in the number of research groups suggesting, on the basis of their studies, that a number of varieties of GM products (mainly maize and soybeans) are as safe and nutritious as the respective conventional non-GM plant, and those raising still serious concerns, was observed. Moreover, it is worth mentioning that most of the studies demonstrating that GM foods are as nutritional and safe as those obtained by conventional breeding, have been performed by biotechnology companies or associates, which are also responsible of commercializing these GM plants. Anyhow, this represents a notable advance in comparison with the lack of studies published in recent years in scientific journals by those companies.
Krimsky S (2015). "An Illusory Consensus behind GMO Health Assessment" (PDF). Science, Technology, & Human Values. 40 (6): 883–914. doi:10.1177/0162243915598381. S2CID 40855100. Archived from the original (PDF) on 7 February 2016. Retrieved 30 October 2016.I began this article with the testimonials from respected scientists that there is literally no scientific controversy over the health effects of GMOs. My investigation into the scientific literature tells another story.
And contrast: Panchin AY, Tuzhikov AI (March 2017). "Published GMO studies find no evidence of harm when corrected for multiple comparisons". Critical Reviews in Biotechnology. 37 (2): 213–217. doi:10.3109/07388551.2015.1130684. PMID 26767435. S2CID 11786594.Here, we show that a number of articles some of which have strongly and negatively influenced the public opinion on GM crops and even provoked political actions, such as GMO embargo, share common flaws in the statistical evaluation of the data. Having accounted for these flaws, we conclude that the data presented in these articles does not provide any substantial evidence of GMO harm. The presented articles suggesting possible harm of GMOs received high public attention. However, despite their claims, they actually weaken the evidence for the harm and lack of substantial equivalency of studied GMOs. We emphasize that with over 1783 published articles on GMOs over the last 10 years it is expected that some of them should have reported undesired differences between GMOs and conventional crops even if no such differences exist in reality.
and Yang YT, Chen B (April 2016). "Governing GMOs in the USA: science, law and public health". Journal of the Science of Food and Agriculture. 96 (6): 1851–5. doi:10.1002/jsfa.7523. PMID 26536836.It is therefore not surprising that efforts to require labeling and to ban GMOs have been a growing political issue in the USA (citing Domingo and Bordonaba, 2011). Overall, a broad scientific consensus holds that currently marketed GM food poses no greater risk than conventional food... Major national and international science and medical associations have stated that no adverse human health effects related to GMO food have been reported or substantiated in peer-reviewed literature to date. Despite various concerns, today, the American Association for the Advancement of Science, the World Health Organization, and many independent international science organizations agree that GMOs are just as safe as other foods. Compared with conventional breeding techniques, genetic engineering is far more precise and, in most cases, less likely to create an unexpected outcome.
- ^ "Statement by the AAAS Board of Directors on Labeling of Genetically Modified Foods" (PDF). American Association for the Advancement of Science. 20 October 2012. Retrieved 8 February 2016.
The EU, for example, has invested more than €300 million in research on the biosafety of GMOs. Its recent report states: "The main conclusion to be drawn from the efforts of more than 130 research projects, covering a period of more than 25 years of research and involving more than 500 independent research groups, is that biotechnology, and in particular GMOs, are not per se more risky than e.g. conventional plant breeding technologies." The World Health Organization, the American Medical Association, the U.S. National Academy of Sciences, the British Royal Society, and every other respected organization that has examined the evidence has come to the same conclusion: consuming foods containing ingredients derived from GM crops is no riskier than consuming the same foods containing ingredients from crop plants modified by conventional plant improvement techniques.
Pinholster G (25 October 2012). "AAAS Board of Directors: Legally Mandating GM Food Labels Could "Mislead and Falsely Alarm Consumers"". American Association for the Advancement of Science. Retrieved 8 February 2016. - ^ European Commission. Directorate-General for Research (2010). A decade of EU-funded GMO research (2001–2010) (PDF). Directorate-General for Research and Innovation. Biotechnologies, Agriculture, Food. European Commission, European Union. doi:10.2777/97784. ISBN 978-92-79-16344-9. Retrieved 8 February 2016.
- ^ "AMA Report on Genetically Modified Crops and Foods (online summary)". American Medical Association. January 2001. Retrieved 19 March 2016.
A report issued by the scientific council of the American Medical Association (AMA) says that no long-term health effects have been detected from the use of transgenic crops and genetically modified foods, and that these foods are substantially equivalent to their conventional counterparts. (from online summary prepared by ISAAA)" "Crops and foods produced using recombinant DNA techniques have been available for fewer than 10 years and no long-term effects have been detected to date. These foods are substantially equivalent to their conventional counterparts. (from original report by AMA: [3])
"Report 2 of the Council on Science and Public Health (A-12): Labeling of Bioengineered Foods" (PDF). American Medical Association. 2012. Archived from the original on 7 September 2012. Retrieved 19 March 2016.Bioengineered foods have been consumed for close to 20 years, and during that time, no overt consequences on human health have been reported and/or substantiated in the peer-reviewed literature.
CS1 maint: bot: original URL status unknown (link) - ^ "Restrictions on Genetically Modified Organisms: United States. Public and Scholarly Opinion". Library of Congress. 9 June 2015. Retrieved 8 February 2016.
Several scientific organizations in the US have issued studies or statements regarding the safety of GMOs indicating that there is no evidence that GMOs present unique safety risks compared to conventionally bred products. These include the National Research Council, the American Association for the Advancement of Science, and the American Medical Association. Groups in the US opposed to GMOs include some environmental organizations, organic farming organizations, and consumer organizations. A substantial number of legal academics have criticized the US's approach to regulating GMOs.
- ^ National Academies of Sciences, Engineering; Division on Earth Life Studies; Board on Agriculture Natural Resources; Committee on Genetically Engineered Crops: Past Experience Future Prospects (2016). Genetically Engineered Crops: Experiences and Prospects. The National Academies of Sciences, Engineering, and Medicine (US). p. 149. doi:10.17226/23395. ISBN 978-0-309-43738-7. PMID 28230933. Retrieved 19 May 2016.
Overall finding on purported adverse effects on human health of foods derived from GE crops: On the basis of detailed examination of comparisons of currently commercialized GE with non-GE foods in compositional analysis, acute and chronic animal toxicity tests, long-term data on health of livestock fed GE foods, and human epidemiological data, the committee found no differences that implicate a higher risk to human health from GE foods than from their non-GE counterparts.
- ^ "Frequently asked questions on genetically modified foods". World Health Organization. Retrieved 8 February 2016.
Different GM organisms include different genes inserted in different ways. This means that individual GM foods and their safety should be assessed on a case-by-case basis and that it is not possible to make general statements on the safety of all GM foods. GM foods currently available on the international market have passed safety assessments and are not likely to present risks for human health. In addition, no effects on human health have been shown as a result of the consumption of such foods by the general population in the countries where they have been approved. Continuous application of safety assessments based on the Codex Alimentarius principles and, where appropriate, adequate post market monitoring, should form the basis for ensuring the safety of GM foods.
- ^ Haslberger AG (July 2003). "Codex guidelines for GM foods include the analysis of unintended effects". Nature Biotechnology. 21 (7): 739–41. doi:10.1038/nbt0703-739. PMID 12833088. S2CID 2533628.
These principles dictate a case-by-case premarket assessment that includes an evaluation of both direct and unintended effects.
- ^ Some medical organizations, including the British Medical Association, advocate further caution based upon the precautionary principle: "Genetically modified foods and health: a second interim statement" (PDF). British Medical Association. March 2004. Retrieved 21 March 2016.
In our view, the potential for GM foods to cause harmful health effects is very small and many of the concerns expressed apply with equal vigour to conventionally derived foods. However, safety concerns cannot, as yet, be dismissed completely on the basis of information currently available. When seeking to optimise the balance between benefits and risks, it is prudent to err on the side of caution and, above all, learn from accumulating knowledge and experience. Any new technology such as genetic modification must be examined for possible benefits and risks to human health and the environment. As with all novel foods, safety assessments in relation to GM foods must be made on a case-by-case basis. Members of the GM jury project were briefed on various aspects of genetic modification by a diverse group of acknowledged experts in the relevant subjects. The GM jury reached the conclusion that the sale of GM foods currently available should be halted and the moratorium on commercial growth of GM crops should be continued. These conclusions were based on the precautionary principle and lack of evidence of any benefit. The Jury expressed concern over the impact of GM crops on farming, the environment, food safety and other potential health effects. The Royal Society review (2002) concluded that the risks to human health associated with the use of specific viral DNA sequences in GM plants are negligible, and while calling for caution in the introduction of potential allergens into food crops, stressed the absence of evidence that commercially available GM foods cause clinical allergic manifestations. The BMA shares the view that there is no robust evidence to prove that GM foods are unsafe but we endorse the call for further research and surveillance to provide convincing evidence of safety and benefit.
- ^ Funk C, Rainie L (29 January 2015). "Public and Scientists' Views on Science and Society". Pew Research Center. Retrieved 24 February 2016.
The largest differences between the public and the AAAS scientists are found in beliefs about the safety of eating genetically modified (GM) foods. Nearly nine-in-ten (88%) scientists say it is generally safe to eat GM foods compared with 37% of the general public, a difference of 51 percentage points.
- ^ Marris C (July 2001). "Public views on GMOs: deconstructing the myths. Stakeholders in the GMO debate often describe public opinion as irrational. But do they really understand the public?". EMBO Reports. 2 (7): 545–8. doi:10.1093/embo-reports/kve142. PMC 1083956. PMID 11463731.
- ^ Final Report of the PABE research project (December 2001). "Public Perceptions of Agricultural Biotechnologies in Europe". Commission of European Communities. Retrieved 24 February 2016.
- ^ Scott SE, Inbar Y, Rozin P (May 2016). "Evidence for Absolute Moral Opposition to Genetically Modified Food in the United States". Perspectives on Psychological Science. 11 (3): 315–324. doi:10.1177/1745691615621275. PMID 27217243. S2CID 261060.
- ^ a b "Genetic Engineering". The Encyclopedia of Science Fiction. 15 May 2017. Retrieved 19 July 2018.
- ^ Koboldt D (29 August 2017). "The Science of Sci-Fi: How Science Fiction Predicted the Future of Genetics". Outer Places. Archived from the original on 19 July 2018. Retrieved 19 July 2018.
- ^ Moraga R (November 2009). "Modern Genetics in the World of Fiction". Clarkesworld Magazine (38). Archived from the original on 19 July 2018.
- ^ a b c Clark M. "Genetic themes in fiction films: Genetics meets Hollywood". The Wellcome Trust. Archived from the original on 18 May 2012. Retrieved 19 July 2018.
Otras lecturas
- British Medical Association (1999). The Impact of Genetic Modification on Agriculture, Food and Health. BMJ Books. ISBN 0-7279-1431-6.
- Donnellan, Craig (2004). Genetic Modification (Issues). Independence Educational Publishers. ISBN 1-86168-288-3.
- Morgan S (1 January 2009). Superfoods: Genetic Modification of Foods. Heinemann Library. ISBN 978-1-4329-2455-3.
- Smiley, Sophie (2005). Genetic Modification: Study Guide (Exploring the Issues). Independence Educational Publishers. ISBN 1-86168-307-3.
- Watson JD (2007). Recombinant DNA: Genes and Genomes: A Short Course. San Francisco: W.H. Freeman. ISBN 978-0-7167-2866-5.
- Weaver S, Michael M (2003). "An Annotated Bibliography of Scientific Publications on the Risks Associated with Genetic Modification". Wellington, NZ: Victoria University. Cite journal requires
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(help) - Zaid A, Hughes HG, Porceddu E, Nicholas F (2001). Glossary of Biotechnology for Food and Agriculture – A Revised and Augmented Edition of the Glossary of Biotechnology and Genetic Engineering. Rome, Italy: FAO. ISBN 92-5-104683-2.
enlaces externos
- GMO Safety - Information about research projects on the biological safety of genetically modified plants.
- GMO-compass, news on GMO en EU