<?xml version="1.0" encoding="ISO-8859-1"?><article xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance">
<front>
<journal-meta>
<journal-id>0122-9761</journal-id>
<journal-title><![CDATA[Boletín de Investigaciones Marinas y Costeras - INVEMAR]]></journal-title>
<abbrev-journal-title><![CDATA[Bol. Invest. Mar. Cost.]]></abbrev-journal-title>
<issn>0122-9761</issn>
<publisher>
<publisher-name><![CDATA[INSTITUTO DE INVESTIGACIONES MARINAS Y COSTERAS "JOSE BENITO VIVES DE ANDRÉIS" (INVEMAR)    INSTITUTO DE INVESTIGACIONES MARINAS Y COSTERAS -JOSE BENITO VIVES DE ANDRÉIS- (INVEMAR)]]></publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id>S0122-97612009000200009</article-id>
<title-group>
<article-title xml:lang="es"><![CDATA[PROBIÓTICOS COMO HERRAMIENTA BIOTECNOLÓGICA EN EL CULTIVO DE CAMARÓN: RESEÑA]]></article-title>
<article-title xml:lang="en"><![CDATA[PROBIOTICS AS A BIOTECHNOLOGICAL TOOL IN SHRIMP CULTURE: A REVIEW]]></article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author">
<name>
<surname><![CDATA[Villamil Díaz]]></surname>
<given-names><![CDATA[Luisa]]></given-names>
</name>
<xref ref-type="aff" rid="A01"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname><![CDATA[Martínez-Silva]]></surname>
<given-names><![CDATA[María Angélica]]></given-names>
</name>
<xref ref-type="aff" rid="A01"/>
</contrib>
</contrib-group>
<aff id="A01">
<institution><![CDATA[,Universidad de Bogotá Jorge Tadeo Lozano Grupo de Investigación en Cultivo y Manejo de Organismos Acuáticos ]]></institution>
<addr-line><![CDATA[Santa Marta ]]></addr-line>
<country>Colombia</country>
</aff>
<pub-date pub-type="pub">
<day>00</day>
<month>12</month>
<year>2009</year>
</pub-date>
<pub-date pub-type="epub">
<day>00</day>
<month>12</month>
<year>2009</year>
</pub-date>
<volume>38</volume>
<numero>2</numero>
<fpage>165</fpage>
<lpage>187</lpage>
<copyright-statement/>
<copyright-year/>
<self-uri xlink:href="http://www.scielo.org.co/scielo.php?script=sci_arttext&amp;pid=S0122-97612009000200009&amp;lng=en&amp;nrm=iso"></self-uri><self-uri xlink:href="http://www.scielo.org.co/scielo.php?script=sci_abstract&amp;pid=S0122-97612009000200009&amp;lng=en&amp;nrm=iso"></self-uri><self-uri xlink:href="http://www.scielo.org.co/scielo.php?script=sci_pdf&amp;pid=S0122-97612009000200009&amp;lng=en&amp;nrm=iso"></self-uri><abstract abstract-type="short" xml:lang="es"><p><![CDATA[Las bacterias probióticas se definen como microorganismos vivos que administrados como suplemento en la dieta pueden causar modificaciones en la microbiota asociada al tracto gastrointestinal del hospedador y generar efectos benéficos como el incremento en la conversión alimentaria, en la resistencia a enfermedades y de la calidad del agua. Durante la última década, su aplicación en el cultivo de camarón se ha hecho frecuente ya que han surgido varios productos comerciales ideados para este fin. Al mismo tiempo, aunque hay publicados varios artículos científicos en el tema, es evidente que hace falta orientar los estudios para entender los mecanismos de acción de los probióticos, así como para establecer los protocolos de aplicación, teniendo en cuenta factores críticos como etapa de cultivo, densidad de siembra y dosis de administración en relación con los mecanismos de defensa inmunitaria del camarón y presencia de organismos potencialmente patógenos. Este trabajo pretende hacer una reseña de las publicaciones más destacadas en el uso de probióticos en acuicultura, particularmente en el cultivo de camarón, ya que su uso se perfila como una de las alternativas con mejores perspectivas al uso indiscriminado de antibióticos que causa problemas tales como la aparición de cepas bacterianas multiresistentes que pueden alterar los ecosistemas próximos al cultivo e incluso afectar la salud del consumidor.]]></p></abstract>
<abstract abstract-type="short" xml:lang="en"><p><![CDATA[Probiotic bacteria are defined as live microorganisms that administered as a diet supplement can cause modifications in the microbiota associated with the gastrointestinal tract of the host and generate beneficial effects such as an increase in the food intake conversion, disease resistance and water quality. During the last decade, its application in shrimp farming has become frequent since several commercial products designed for this purpose have emerged. At the same time, although there are a number of scientific articles published on the subject, it is evident that there is a lack of studies oriented to understand the probiotics working mechanisms and to establish the protocols for its implementation, taking into account critical factors such as the stage of cultivation, culture density and dosage in relation to the shrimp immune defense mechanisms and presence of potentially pathogenic organisms. This paper aims to review the most prominent publications regarding the use of probiotics in aquaculture, particularly in shrimp farming, since its use seems to be the alternative with better perspectives to the indiscriminate use of antibiotics that cause problems such as the emergence of multi-resistant bacterial strains that could alter the ecosystems near acuaculture sites and even affect consumer health.]]></p></abstract>
<kwd-group>
<kwd lng="es"><![CDATA[Bacteria]]></kwd>
<kwd lng="es"><![CDATA[Cultivo de camarón]]></kwd>
<kwd lng="es"><![CDATA[Probióticos]]></kwd>
<kwd lng="es"><![CDATA[Bacillus]]></kwd>
<kwd lng="es"><![CDATA[Lactobacillus]]></kwd>
<kwd lng="en"><![CDATA[Bacteria]]></kwd>
<kwd lng="en"><![CDATA[Shrimp culture]]></kwd>
<kwd lng="en"><![CDATA[Probiotics]]></kwd>
<kwd lng="en"><![CDATA[Bacillus]]></kwd>
<kwd lng="en"><![CDATA[Lactobacillus]]></kwd>
</kwd-group>
</article-meta>
</front><body><![CDATA[  <font face="verdana" size="2">     <p align="center"><font size="4"><b>PROBI&Oacute;TICOS COMO HERRAMIENTA BIOTECNOL&Oacute;GICA EN EL CULTIVO  DE CAMAR&Oacute;N: RESE&Ntilde;A</b></font></p>     <p align="center"><font size="3"><b>PROBIOTICS AS A BIOTECHNOLOGICAL TOOL IN SHRIMP CULTURE: A REVIEW</b></font></p>     <p>&nbsp;</p>     <p><b>Luisa Villamil D&iacute;az y Mar&iacute;a Ang&eacute;lica Mart&iacute;nez-Silva</b></p>     <p><i>Universidad de Bogot&aacute; Jorge Tadeo Lozano, Grupo de  Investigaci&oacute;n en Cultivo y Manejo de Organismos Acu&aacute;ticos, Santa Marta,  Colombia. <a href="mailto:luisa.villamil@utadeo.edu.co">luisa.villamil@utadeo.edu.co</a> (L.V.D.), <a href="mailto:maria.martinez@utadeo.edu.co">maria.martinez@utadeo.edu.co</a> (M.A.M.S.)</i></p> <hr size="1" />     <p>&nbsp;</p>     <p><b>RESUMEN</b></p>     <p>Las bacterias probi&oacute;ticas se definen como microorganismos vivos que  administrados como suplemento en la dieta pueden causar modificaciones en la  microbiota asociada al tracto gastrointestinal del hospedador y generar efectos  ben&eacute;ficos como el incremento en la conversi&oacute;n alimentaria, en la resistencia a  enfermedades y de la calidad del agua. Durante la &uacute;ltima d&eacute;cada, su aplicaci&oacute;n  en el cultivo de camar&oacute;n se ha hecho frecuente ya que han surgido varios  productos comerciales ideados para este fin. Al mismo tiempo, aunque hay  publicados varios art&iacute;culos cient&iacute;ficos en el tema, es evidente que hace falta  orientar los estudios para entender los mecanismos de acci&oacute;n de los  probi&oacute;ticos, as&iacute; como para establecer los protocolos de aplicaci&oacute;n, teniendo en  cuenta factores cr&iacute;ticos como etapa de cultivo, densidad de siembra y dosis de  administraci&oacute;n en relaci&oacute;n con los mecanismos de defensa inmunitaria del  camar&oacute;n y presencia de organismos potencialmente pat&oacute;genos. Este trabajo  pretende hacer una rese&ntilde;a de las publicaciones m&aacute;s destacadas en el uso de  probi&oacute;ticos en acuicultura, particularmente en el cultivo de camar&oacute;n, ya que su  uso se perfila como una de las alternativas con mejores perspectivas al uso  indiscriminado de antibi&oacute;ticos que causa problemas tales como la aparici&oacute;n de  cepas bacterianas multiresistentes que pueden alterar los ecosistemas pr&oacute;ximos  al cultivo e incluso afectar la salud del consumidor.</p>     <p><i>PALABRAS CLAVE:</i> Bacteria, Cultivo de  camar&oacute;n, Probi&oacute;ticos, <i>Bacillus</i>, <i>Lactobacillus</i>.</p> <hr size="1" />     ]]></body>
<body><![CDATA[<p>&nbsp;</p>     <p><b>ABSTRACT</b></p>     <p>Probiotic bacteria are defined as live  microorganisms that administered as a diet supplement can cause modifications  in the microbiota associated with the gastrointestinal tract of the host and  generate beneficial effects such as an increase in the food intake conversion,  disease resistance and water quality. During the last decade, its application  in shrimp farming has become frequent since several commercial products  designed for this purpose have emerged. At the same time, although there are a  number of scientific articles published on the subject, it is evident that  there is a lack of studies oriented to understand the probiotics working  mechanisms and to establish the protocols for its implementation, taking into  account critical factors such as the stage of cultivation, culture density and  dosage in relation to the shrimp immune defense mechanisms and presence of  potentially pathogenic organisms. This paper aims to review the most prominent  publications regarding the use of probiotics in aquaculture, particularly in  shrimp farming, since its use seems to be the alternative with better  perspectives to the indiscriminate use of antibiotics that cause problems such  as the emergence of multi-resistant bacterial strains that could alter the  ecosystems near acuaculture sites and even affect consumer health.</p>     <p><i>KEY WORDS:</i> Bacteria, Shrimp culture,  Probiotics, <i>Bacillus</i>, <i>Lactobacillus</i>.</p> <hr size="1" />     <p>&nbsp;</p>     <p><b>INTRODUCCI&Oacute;N</b></p>     <p>El dram&aacute;tico incremento en la poblaci&oacute;n  mundial en los &uacute;ltimos dos siglos, sumado a la sobreexplotaci&oacute;n de diferentes  pesquer&iacute;as (FAO, 2006) son factores que explican en parte el d&eacute;ficit de prote&iacute;na  para consumo humano. De acuerdo con el informe de la FAO, Estado de la  Acuicultura Mundial: 2006, la acuicultura es una actividad con un alto  potencial para satisfacer la creciente demanda de alimentos acu&aacute;ticos ya que es  probablemente el sector productivo de m&aacute;s r&aacute;pido crecimiento, que genera  actualmente alrededor del 50% de la producci&oacute;n de pescado en el mundo. El  camar&oacute;n es uno de los productos cultivados m&aacute;s importantes en Asia y la regi&oacute;n  Pac&iacute;fica, con una producci&oacute;n de 1.1 millones de toneladas para 2006, mientras  que Latinoam&eacute;rica alcanz&oacute; 260000 toneladas (FAO, 2006).</p>     <P>Una de las principales  dificultades que existe en el cultivo comercial de organismos marinos es la  aparici&oacute;n de enfermedades infecciosas como resultado de la incidencia de  bacterias, hongos y virus frecuentemente asociados con el aumento en las  densidades de cultivo y r&aacute;pido desarrollo de la acuicultura con deficiencias en  los m&eacute;todos de manejo, calidad de aguas, valor nutricional del alimento, entre  otros factores (Paillard <i>et al</i>., 2004;  Pruzzo <i>et al</i>., 2005). En el  subsector cultivador de camar&oacute;n marino, las enfermedades son consideradas  actualmente como el principal factor causante de perdidas en producci&oacute;n y  dinero (Lin, 1995; Lightner, 1996; Moriarty, 1999). Enfermedades infecciosas  como el virus del Taura, la Necrosis Hematopoy&eacute;tica Hipodermica (IHHN) y el  s&iacute;ndrome de la mancha blanca (WSSV) han generado devastaci&oacute;n en Sur Am&eacute;rica,  Hawai y el Pacifico. A este problema de las enfermedades se suma, adem&aacute;s, el  bajo precio que actualmente tiene el camar&oacute;n en el mercado (US) (FAO, 2006),  por lo que el sector debe buscar estrategias de producci&oacute;n que le aporten mayor  competitividad a la cadena productiva, facilitando obtener una mayor cantidad  de individuos y de mejor calidad por el mismo costo.</P>     <p>En el pasado, ante la aparici&oacute;n de  cualquier s&iacute;ntoma de enfermedad, se recurri&oacute; al uso incontrolado de agentes  qu&iacute;micos como los antibi&oacute;ticos, cuyo espectro de acci&oacute;n es obviamente limitado  para la prevenci&oacute;n y control de enfermedades (Subasinghe, 1997) y, adem&aacute;s,  causan efectos adversos como la aparici&oacute;n de cepas bacterianas  multi-resistentes que incluso pueden llegar a afectar la salud del consumidor  (Tsoumas <i>et al</i>,. 1989; Smith <i>et al</i>., 1994). De igual manera, debido al largo  tiempo de vida media en el agua de algunos antibi&oacute;ticos como la  oxitetraciclina, los residuos de antibi&oacute;ticos generados en granjas de cultivo  en algunos pueden modificar las comunidades microbianas de los ecosistemas  pr&oacute;ximos (Hektoen <i>et al</i>, 1995). De  acuerdo con estos indicios, es claro que los acuicultores necesitan evitar la  aplicaci&oacute;n innecesaria de antibi&oacute;ticos y entender la complejidad de la  comunidad microbiana que est&aacute; presente en el agua de cultivo e implementar la  aplicaci&oacute;n de bacterias ben&eacute;ficas para combatir a las pat&oacute;genas y las  eventuales mortalidades a las que pudiera dar lugar. Considerando el resultado  exitoso que se ha obtenido en algunos experimentos cient&iacute;ficos realizados en granjas  de peces, camarones y ostras, la Organizaci&oacute;n de las Naciones Unidas para la  Alimentaci&oacute;n y la Agricultura, FAO (Subasinghe, 1997), defini&oacute; el desarrollo de  vacunas efectivas, el uso de inmunoestimulantes, potenciadores inmunol&oacute;gicos y  de probi&oacute;ticos para mejorar la calidad del medio acu&aacute;tico, como las principales  &aacute;reas de investigaci&oacute;n en la lucha contra las enfermedades en acuicultura.</p>     <p>En Colombia, a pesar de que actualmente  se usan probi&oacute;ticos comerciales en el cultivo de organismos acu&aacute;ticos, no se  han realizado estudios en los que se eval&uacute;e a profundidad los beneficios de su  aplicaci&oacute;n, as&iacute; como resolver otras preguntas como comparar el efecto de los  probi&oacute;ticos vivos con las c&eacute;lulas inactivas, como lo proponen Villamil <i>et al</i>. (2003b) y Gatesoupe (2008), o probar  otros compuestos como bacteriocinas. Cabe mencionar que los productos de  probi&oacute;ticos que se comercializan en el pa&iacute;s son importados, por lo cual la  realizaci&oacute;n de proyectos encaminados a obtener una formulaci&oacute;n de probi&oacute;ticos  con aislados bacterianos nativos a escala comercial que pueda ser probado con  las caracter&iacute;sticas propias de nuestros cultivos, ser&iacute;a un importante avance en  la prevenci&oacute;n de las enfermedades que afectan dram&aacute;ticamente el sector  acuicultor.</p>     ]]></body>
<body><![CDATA[<p><b>¿QU&Eacute;  ES UN PROBI&Oacute;TICO?</b></p>     <p>El termino probi&oacute;tico  ha sufrido modificaciones en su significado a lo largo de los a&ntilde;os. De esta  manera en 1968, se defini&oacute; como un suplemento microbiano que se suministra a  animales y humanos. Fuller (1989) lo redefini&oacute; como un microorganismo vivo que  se administra al hospedero suplementado en el alimento para beneficiar el  balance microbiano intestinal. Posteriormente, el t&eacute;rmino fue usado para  referirse a un adyuvante dietario microbiano administrado de tal manera que se  mantenga vivo dentro del tracto gastrointestinal, y que beneficie la fisiolog&iacute;a  del hospedero modulando el sistema inmune, as&iacute; como mejorando el balance  microbiano mediante la prevenci&oacute;n de la colonizaci&oacute;n de bacterias indeseables  en el tracto intestinal (Gatesoupe, 1999; Naidu <i>et al</i>, 1999). Verschuere <i>et al</i>. (2000b) dieron una definici&oacute;n m&aacute;s amplia  de los probi&oacute;ticos como microorganismos vivos que tienen efectos ben&eacute;ficos en  el hospedero mediante la modificaci&oacute;n de la microbiota asociada, el incremento  del aprovechamiento de la comida, el mejoramiento de la respuesta a  enfermedades y de la calidad del ambiente. Sin embargo, en este punto es  importante se&ntilde;alar que las bacterias que simplemente cumplen alguno de estos  roles, tales como la producci&oacute;n de nutrientes esenciales para el  aprovechamiento de las especies cultivadas, o bacterias que solamente ejercen  una funci&oacute;n espec&iacute;fica de bio-remediaci&oacute;n en el medio ambiente, no deben  considerarse como probi&oacute;ticos. La aplicaci&oacute;n de probi&oacute;ticos como control biol&oacute;gico  es una alternativa viable dada la habilidad que poseen las cepas seleccionadas  para impedir el crecimiento de bacterias oportunistas e influir en general en  el establecimiento de la comunidad microbiana tanto en los individuos como en  el agua de cultivo.</p>     <p>Aunque la evaluaci&oacute;n de probi&oacute;ticos en  acuicultura se ha abordado en diferentes art&iacute;culos cient&iacute;ficos y tambi&eacute;n est&aacute;n  disponibles en el mercado productos comerciales, el concepto de  &quot;probi&oacute;tico&quot; y su eficacia sigue siendo poco conocido y controvertido;  existen dudas sobre la eficiencia y seguridad de los organismos probi&oacute;ticos que  pueden tener origen en una multiplicidad de factores tales como el uso de  aislados con poca actividad, en que los productos comerciales han generado  expectativas poco realistas por parte de los consumidores, adem&aacute;s porque  relativamente pocos estudios han abordado los mecanismos de acci&oacute;n de la cepas  probi&oacute;ticas seleccionadas utilizando condiciones estandarizadas. Por lo cual,  es urgente para el sector cultivador de camar&oacute;n que se realicen estudios  cient&iacute;icos reproducibles y exactos a escala piloto y comercial para lograr la  aplicaci&oacute;n de un protocolo que garantice una mejora significativa en los niveles  productivos, teniendo aspectos clave como la fase del ciclo productivo, el tiempo,  dosis y v&iacute;a de administraci&oacute;n, entre otros factores. De igual manera, los  procesos de selecci&oacute;n, crecimiento y escalamiento en la producci&oacute;n de los  microorganismos probi&oacute;ticos seleccionados debe realizarse con alta rigurosidad  de tal manera que cumplan los est&aacute;ndares de calidad, ya que podr&iacute;a alterarse la  composici&oacute;n de las mezclas bacterianas originales, con resultados  impredecibles.</p>     <p><b>¿CU&Aacute;LES SON LOS  MECANISMOS DE ACCI&Oacute;N DE LOS PROBI&Oacute;TICOS?</b></p>     <p>Las publicaciones  cient&iacute;ficas existentes en el tema han facilitado el entendimiento de los modos  de acci&oacute;n de los probi&oacute;ticos en el hospedero, entre ellos la competencia por  nutrientes, la modulaci&oacute;n de la respuesta inmunitaria no espec&iacute;ica, la  producci&oacute;n de compuestos antimicrobianos, la competencia por el sitio de  ijaci&oacute;n en el tracto gastrointestinal, entre otros que se han evidenciado en  experimentos <i>in vitro</i> e  <i>in vivo</i>. Sin embargo, es necesario precisar que la  eficacia de un probi&oacute;tico seleccionado <i>in vitro</i> puede  cambiar cuando se administra al hospedero, dada la gran influencia de factores  m&aacute;s complejos como la ingesti&oacute;n selectiva (Prieur, 1981; Riquelme <i>et al</i>, 2000) y la muerte en el tracto  gastrointestinal (Vine <i>et al</i>, 2006)  causada por la incapacidad del probi&oacute;tico para mantener su fisiolog&iacute;a bajo  circunstancias de una mayor interacci&oacute;n microbiana (Tinh <i>et al</i>, 2007). En general, la relaci&oacute;n entre los  experimentos <i>in vivo</i> e <i>in vitro</i> ha sido descrita en los &uacute;ltimos art&iacute;culos  de revisi&oacute;n cient&iacute;fica de uso de probi&oacute;ticos en acuicultura (Manto y Austin,  2002; Balcazar <i>et al</i>, 2006; Vine <i>et al</i>, 2006; Tinh <i>et al</i>, 2007; Gatesoupe, 2008). Entre los  principales mecanismos descritos que usan los aislados probi&oacute;ticos para  beneficiar al hospedador, se encuentran:</p>     <p><b>a. Colonizaci&oacute;n y adhesi&oacute;n en el tracto  gastrointestinal</b></p>     <p>Es bien sabido que la habilidad de las  bacterias para adherirse y sobrevivir en el mucus ent&eacute;rico es decisiva en el  establecimiento de la microbiota intestinal. La capacidad de adherencia es una  caracter&iacute;stica que es aprovechada de igual manera tanto por las bacterias  probi&oacute;ticas como por las pat&oacute;genas. En el caso de las probi&oacute;ticas, &eacute;ste ha sido  uno de los criterios m&aacute;s importantes para su selecci&oacute;n y aplicaci&oacute;n en  acuicultura (Salminen <i>et al</i>, 1996;  Nikoskelainen <i>et al</i>, 2003), mientras  que para las pat&oacute;genas la habilidad para adherirse, se relaciona con la  virulencia y se considera como el primer paso para una infecci&oacute;n (Bengmark,  1998). En acuicultura, la informaci&oacute;n disponible indica que las bacterias  aisladas de animales cultivados o de su entorno tienen mayor capacidad de  adhesi&oacute;n al mucus gastrointestinal y a los tejidos, que las de otras bacterias  for&aacute;neas que suelen ser transitorias, por lo que surge la necesidad de que los  probi&oacute;ticos sean continuamente administrados, ya sea como suplemento en el  alimento o a trav&eacute;s del agua de cultivo (Ring&oslash; y Gatesoupe, 1998; Villamil <i>et al</i>, 2003c). Adem&aacute;s, se ha documentado que  aislados microbianos de un organismo pueden colonizar otras especies  cultivadas, indicando as&iacute; la falta de especificidad para la colonizaci&oacute;n del  tracto digestivo (Ring&oslash;, 1999).</p>     <p><b>b. Producci&oacute;n de antibi&oacute;ticos / compuestos  antivirales</b></p>     <p>La selecci&oacute;n de microorganismos con  actividad probi&oacute;tica tambi&eacute;n se puede determinar por la capacidad de generar  productos extracelulares (ECPS) que pueden inhibir o matar otras bacterias  potencialmente pat&oacute;genas, entre ellos sustancias antibacteriales (Imada <i>et al</i>, 1985; Williams y Vickers, 1986; Maeda,  1994; V&aacute;zquez <i>et al</i>, 2006,),  sider&oacute;foros (Spanggaard <i>et al</i>, 2001;  Brunt <i>et al</i>, 2007), enzimas  bacteriol&iacute;ticas (Nair <i>et al</i>, 1985), &aacute;cido  l&aacute;ctico (Lindgren y Clevstrom, 1978; Alakomi <i>et al</i>, 2000), &aacute;cidos org&aacute;nicos (Midolo <i>et al</i>, 1995), per&oacute;xido de hidrogeno (V&aacute;zquez <i>et al</i>., 2005), di&oacute;xido de carbono (Naidu <i>et al</i>., 1999) y bacteriocinas (Villamil <i>et al</i>., 2003a;  Gatesoupe 2008). Las bacteriocinas son polip&eacute;ptidos  bacteriost&aacute;ticos o bactericidas que en su mayor&iacute;a son activos contra bacterias  estrechamente relacionadas (Klaenhammer,  1998) y microorganismos Gram-positivos (Hurst, 1981; Severina <i>et al</i>., 1998). Una de las bacteriocinas mejor conocidas es la nisina, que es un  p&eacute;ptido sintetizado ribosomalmente y producido por algunas cepas de Lactococcus lactis. La eficiencia de este p&eacute;ptido ha sido  probada contra pat&oacute;genos multiresistentes de humanos como <i>Streptococcus pneumoniae</i>, <i>Staphylococcus aureus</i>, <i>Staphylococcus epidermidis</i> y <i>Enterococcus faecalis</i>, entre otros (Hurst,  1981; Liu y Hansen, 1990).</p>     <p>Por otra parte, los  p&eacute;ptidos antimicrobianos derivados de estas bacterias interact&uacute;an con las  c&eacute;lulas del sistema inmunol&oacute;gico (De Pablo <i>et al</i>., 1999), y pueden ser reconocidos por ellas,  gracias a los anticuerpos policlonales contra bacteriocinas como la nisina y  pediocina en peces. Adicionalmente, algunas bacterias acido-l&aacute;cticas (LAB)  tambi&eacute;n poseen actividad contra bacterias Gram-negativas como <i>Vibrio anguillarum</i>, <i>Vibrio salmonicida</i> y <i>Proteus vulgaris</i> (Ring&oslash; y Gatesoupe, 1998). La carnocina de <i>Carnobacterium piscicola</i> tambi&eacute;n ha resultado eficaz para combatir  a <i>Aeromonas  hydrophila</i> (Lewus  <i>et al</i>., 1991). Estudios m&aacute;s recientes han postulado un  nuevo g&eacute;nero de bacterias aisladas de granjas productoras de rodaballo (<i>Scophtahlmus maximus</i>), Roseobacter, como probi&oacute;ticos basado en la actividad  antibacteriana contra bacterias pat&oacute;genas de peces marinos <i>V. anguillarum</i> y <i>V. splendidus</i> (Hjelm <i>et al</i>., 2004).</p>     ]]></body>
<body><![CDATA[<p>Los probi&oacute;ticos no solo tienen capacidad  antibacteriana, tambi&eacute;n se ha descrito actividad antiviral de algunos aislados  como <i>Pseudomonas sp</i>., <i>Vibrio spp</i>. y <i>Aeromonas sp</i>., contra el virus de la necrosis  hematopoy&eacute;tica (IHNV) (Kamei <i>et al</i>., 1988). Maeda <i>et al</i>. (1997) aislaron una cepa de <i>Pseudoalteromonas undina</i>, que ejerci&oacute; efectos antivirales  e increment&oacute; la  supervivencia en camar&oacute;n (<i>Penaeus sp</i>.)  y dorada (<i>Sparus aurata</i>) infectados  experimentalmente con el virus de la necrosis neuro Sima-aji (SJNNV), Baculo  e Iridovirus.  Varias cepas de Vibrio  aisladas de un cultivo  de camar&oacute;n, tambi&eacute;n presentaron actividad antiviral significativa especialmente  frente a IHNV y el virus de <i>Oncorhynchus masou</i> (OMV) (Direkbusarakom <i>et al</i>. 1998), aunque estudios recientes est&aacute;n dirigidos a establecer el mecanismo  de actividad antiviral de manera directa en la supervivencia evaluando otros  factores m&aacute;s complejos. Otros mecanismos que podr&iacute;an inhibir el crecimiento de  bacterias indeseables, como la competencia por nutrientes, la energ&iacute;a  disponible o sitios de adherencia (Gatesoupe, 1999; Verschuere <i>et al</i>., 2000a) deben ser tenidos en cuenta.</p>     <p><b>c. Producci&oacute;n de compuestos ben&eacute;ficos</b></p>     <p>Las bacterias marinas y las levaduras  pueden llegar a ser un recurso de prote&iacute;na importante en el mejoramiento del  aporte nutricional de algunas especies acu&aacute;ticas cultivadas debido al perfil de  amino&aacute;cidos que contienen (Brown <i>et al</i>, 1996).  En ciertos aislados de bacterias intestinales, se ha demostrado alta producci&oacute;n  de &aacute;cidos grasos de cadena corta (Yazawa, 1996) y tambi&eacute;n su contribuci&oacute;n al  valor nutritivo de los rot&iacute;feros (Watanabe <i>et al</i>, 1992) y peces (Clements, 1997). De la  misma manera, los l&iacute;pidos producidos por microorganismos marinos han sido  descritos como sustancias de gran importancia para la nutrici&oacute;n de especies  acu&aacute;ticas como el rodaballo y la tilapia (Ring&oslash; <i>et al</i>, 1992; Kihara y Sakata, 1997). Por otra  parte, la producci&oacute;n de enzimas como lipasas, quitinasas y proteasas, por parte  de microorganismos seleccionados, pueden contribuir al proceso digestivo de los  organismos cultivados, especialmente en estadios larvales de bivalvos (Prieur <i>et al</i>, 1990) y camar&oacute;n (Wang <i>et al</i>, 2000).</p>     <p><b>d. Mejoramiento de las funciones inmunes</b></p>     <p>A pesar de que existe un amplio n&uacute;mero de  publicaciones cient&iacute;ficas en las que se describe un aumento en la resistencia  de peces tratados con probi&oacute;ticos durante infecciones experimentales  (Gatesoupe, 1994; Cai <i>et al</i>, 1998, Ottessen y  Olafsen, 2000; Robertson <i>et al</i>, 2000;  Balc&aacute;zar <i>et al</i>, 2006), existen  relativamente muy pocas en las que se estudien a fondo los mecanismos empleados  en dicha defensa; s&oacute;lo trabajos recientes han demostrado la incidencia de los  probi&oacute;ticos en las funciones del sistema inmune. Manto y Austin (2002)  describieron un incremento en par&aacute;metros celulares, como el n&uacute;mero de  eritrocitos, linfocitos y macr&oacute;fagos y un aumento de la actividad lisoz&iacute;mica de  <i>Salmo salar</i>,  <i>Oncorhynchus mykiss</i> y <i>Scophthalmus maximus</i> alimentados con probi&oacute;ticos  seleccionados, tanto Gram-positivos como Gram negativos. Villamil <i>et al</i>. (2002) evaluaron los efectos  inmunomoduladores de varias cepas de LAB de origen terrestre, encontrando que <i>L. lactis</i> viable e inactivado por calor incrementa  funciones inmunitarias de rodaballo (<i>S. maximus</i>), como quimioluminiscencia de macr&oacute;fagos de ri&ntilde;on anterior y  concentraci&oacute;n de lisozima en suero. M&aacute;s tarde, Villamil <i>et al</i>. (2003b) encontraron que, en el caso de  LAB, no s&oacute;lo las c&eacute;lulas enteras son capaces de inducir un aumento en la  respuesta inmune, algunos productos extracelulares como la nisina, principal  bacteriocina producida por <i>L. lactis</i>, pueden  aumentar la quimioluminiscencia y la producci&oacute;n de &oacute;xido n&iacute;trico en una dosis y  tiempo dependiente en rodaballo (<i>S. maximus</i>).</p>     <p>En camar&oacute;n, Balc&aacute;zar (2003) describi&oacute; un  aumento en la resistencia de <i>Litopenaeus vannamei</i>, alimentado con un suplemento de <i>Bacillus</i> y  <i>Vibrio</i>, contra <i>Vibrio harveyi</i> y el s&iacute;ndrome de mancha blanca, este  incremento en la resistencia se correlacion&oacute; con un aumento de la fagocitosis y  la actividad antibacteriana de los hemocitos. Chiu <i>et al</i>. (2007) informaron que el camar&oacute;n blanco <i>L. vannamei</i> tratado con complemento alimenticio de <i>Lactobacillus plantarum</i> aument&oacute; significativamente la actividad  fenoloxidasa (PO), el estallido respiratorio y la super&oacute;xido dismutasa (SOD), as&iacute; como la transcripci&oacute;n del mRNA de  peroxinectina (PE) y profenoxidasa (proPO), lo que contribuy&oacute; a la eliminaci&oacute;n  de <i>Vibrio  alginolyticus</i> durante  infecciones experimentales.</p>     <p><b>e. Mejora de la calidad de agua</b></p>     <p>Se ha propuesto que las  bacterias del g&eacute;nero <i>Bacillus</i> seleccionadas como probi&oacute;ticos pueden convertir la materia org&aacute;nica en  CO<sub>2</sub>, en contraste con las bacterias Gram-negativas que se  caracterizan por convertir materia org&aacute;nica en biomasa bacteriana o limo  (Dalmin <i>et al</i>., 2001). Laloo <i>et al</i>. (2007) comprobaron la capacidad de tres aislados del genero <i>Bacillus</i> para diminuir las concentraciones de  nitritos, nitratos y amonios en el agua de cultivo de peces ornamentales. Este  mismo fen&oacute;meno tambi&eacute;n fue observado por Kim <i>et al</i>. (2005) en <i>B.  subtilus</i>, <i>B. cereus</i> y  <i>B. licheniformis</i>, quienes atribuyen estos efectos a  mecanismos tales como bioacumulaci&oacute;n, bio-asimilaci&oacute;n y nitrificaci&oacute;n. Aunque  la eliminaci&oacute;n de nitr&oacute;geno es una propiedad predominante en bacterias  autotr&oacute;icas, se han producido varios informes que sugieren una contribuci&oacute;n de  las bacterias heter&oacute;trofas en este sentido (Abou-Seada y Ottow,  1985; Robertson  y Kuenen, 1990; Sakai <i>et al</i>., 1996, 1997; Kim <i>et al</i>., 2005; Lin <i>et al</i>., 2006). De manera  controversial,  hay publicados varios estudios en camar&oacute;n y bagre que no pudieron confirmar  &eacute;stas hip&oacute;tesis (Queiroz y Boyd, 1998; Rengpipat <i>et al</i>., 1998). Adicionalmente, la interacci&oacute;n entre bacterias probi&oacute;ticas y  microalgas en los tanques de cultivo en general produce efectos positivos, ya  que estabiliza los factores nutricionales del alimento vivo pudiendo contribuir  al establecimiento de la microlora intestinal beneiciosa de los hospederos  (Reitan <i>et al</i>., 1993, 1997).</p>     <p><b>USO DE PROBI&Oacute;TICOS EN CAMAR&Oacute;N Y OTROS CULTIVOS</b></p>     <p>La mayor&iacute;a de los  microorganismos probi&oacute;ticos propuestos para acuicultura pertenecen a las  bacterias &aacute;cido-l&aacute;cticas (LAB), de los cuales los g&eacute;neros m&aacute;s utilizados son  <i>Lactobacillus</i>  y <i>Lactococcus</i>. Estos son considerados como GRAS  (&quot;Generally recognized as safe&quot;), reduciendo de este modo la necesidad de ensayos de  seguridad biol&oacute;gica, inevitables para garantizar que la implementaci&oacute;n de  aislados probi&oacute;ticos no va a causar da&ntilde;os colaterales a los organismos  cultivados ni al consumidor inal (Holzapfel <i>et al</i>, 1998). El uso de probi&oacute;ticos como LAB  est&aacute; relativamente bien establecido en otras especies animales (Wallace y  Newbold, 1992; Aiba <i>et al</i>, 1998; Kontula <i>et al</i>, 1998; Kirjavainen <i>et al</i>, 1999a, 1999b; Netherwood <i>et al</i>, 1999), de ellos se destaca el aumento de  tama&ntilde;o y peso, el establecimiento de un equilibrio microbiano intestinal, as&iacute;  como la mejora de algunas respuestas inmunes.</p>     ]]></body>
<body><![CDATA[<p>En peces, las LAB se han descrito como  parte de la microflora normal de los organismos (Str&oslash;m y Olfasen, 1990, Ring&oslash; y  Str&oslash;m, 1994; Ring&oslash; <i>et al</i>, 1998; Robertson <i>et al</i>, 2000). La administraci&oacute;n de LAB ex&oacute;genas  tambi&eacute;n se ha asociado con la inhibici&oacute;n del crecimiento de bacterias pat&oacute;genas  (Lewus <i>et al</i>, 1991; Gildberg <i>et al</i>, 1995; Santos <i>et al</i>, 1996), como promotor del crecimiento de  peces (Noh <i>et al</i>, 1994) y, en  algunos casos, con un aumento de la supervivencia de peces infectados  experimentalmente (Gatesoupe, 1994; Gildberg, 1995; Robertson <i>et al</i>, 2000). La prevenci&oacute;n de la colonizaci&oacute;n  de bacterias perjudiciales con una selecci&oacute;n de cepas bacterianas, se ha  propuesto como una alternativa importante para el control microbiano en el  cultivo de <i>Artemia</i>  (Verschuere <i>et al</i>, 1999). Tambi&eacute;n se ha demostrado que  algunas de &eacute;stas cepas bacterianas seleccionadas pueden prevenir el crecimiento  de bacterias pat&oacute;genas como <i>Vibrio proteolyticus</i> en el cultivo de <i>Artemia</i> (Verschuere  <i>et al</i>. ,1999). Recientemente, Balc&aacute;zar <i>et al</i>. (2007a, 2007b, 2007c) demostraron que <i>Lactococcus lactis ssp</i>. <i>lactis</i> y  <i>Leuconostoc mesenteroides</i> aislados de salm&oacute;nidos, eran capaces de persistir en el intestino de  la trucha arco iris (<i>Salmo  trutta</i>) y aumentar  significativamente la actividad de la lisozima despu&eacute;s de la suplementaci&oacute;n de  alimentos con probi&oacute;ticos.</p>     <p>Las LAB tambi&eacute;n se han aplicado con &eacute;xito  en el cultivo de rot&iacute;feros (<i>Brachionus plicatilis</i>). Gatesoupe (1991) inform&oacute; que un preparado comercial de <i>Lactobacillus plantarum</i> en estado viable disminuy&oacute; el recuento de  <i>A. salmonicida</i> y otras bacterias asociadas a los  rot&iacute;feros. Harzevili <i>et al</i>. (1998)  demostraron que <i>L.  lactis</i> (AR21)  tienen un efecto inhibitorio contra <i>V. anguillarum</i> en cultivo de rot&iacute;feros en condiciones sub&oacute;ptimas y, de igual forma,  demuestra un incremento significativo en el crecimiento en condiciones de  alimentaci&oacute;n &oacute;ptimas.</p>     <p>En general, el uso de probi&oacute;ticos en el  cultivo de camar&oacute;n ha tenido buenas perspectivas; diversas publicaciones  cient&iacute;ficas han demostrado efectos positivos de la aplicaci&oacute;n de probi&oacute;ticos  como se muestra en la <a href="#tab1">Tabla 1</a>. Sin embargo, los resultados obtenidos en algunas  granjas pueden ser variables debido a diferentes factores, que pueden afectar  el resultado de las operaciones a largo plazo, como la calidad del probi&oacute;tico,  el modo de administraci&oacute;n, la talla y las especies de camar&oacute;n.</p>     <p align="center"><img src="img/revistas/mar/v38n2/v38n2a09tab1.gif"><a name="tab1"></a></p>     <p>Como se muestra en la  <a href="#tab1">Tabla 1</a>, los mayores beneficios obtenidos por la suplementaci&oacute;n de probi&oacute;ticos  en el cultivo de crust&aacute;ceos son el incremento de la supervivencia durante  infecciones experimentales, presumiblemente asociado con la potenciaci&oacute;n de las  defensas del sistema inmune innato, as&iacute; como un efecto antimicrobiano directo  de los asilados demostrado en varios estudios, especialmente in vitro. Esta condici&oacute;n est&aacute; tambi&eacute;n relacionada  con cambios significativos en la microbiota intestinal, por el desplazamiento  de bacterias oportunistas y pat&oacute;genas y al mismo tiempo con la colonizaci&oacute;n  dominante de las cepas de microorganismos usados como probi&oacute;ticos.</p>     <p>Otro efecto ben&eacute;fico asociado al uso de  probi&oacute;ticos es el aumento de la tasa de crecimiento y de incremento en peso,  que es atribuido principalmente al establecimiento en el tracto  gastrointestinal (TGI) del hospedero, contribuyendo al balance de la microflora  y mejorando la absorci&oacute;n de nutrientes (Gatesoupe, 2008) debido a la producci&oacute;n  de enzimas digestivas (proteasas y amilasas) y exoenzimas cuya funci&oacute;n es  romper la celulosa y el almid&oacute;n del alimento, facilitando de esta manera su  asimilaci&oacute;n (Jory, 1998). Como una acci&oacute;n positiva adicional tambi&eacute;n  documentada en acuicultura, es importante mencionar la mejora en la calidad del  agua relacionada con la capacidad de algunas bacterias para reducir el amonio y  la materia org&aacute;nica que son frecuentemente una de las causas de estr&eacute;s en los  cultivos de camar&oacute;n. Sin embargo, de acuerdo con la estricta definici&oacute;n de  probi&oacute;tico, es importante que quede claro que los microorganismos que son  capaces de mejorar la calidad del agua, sin causar ning&uacute;n efecto positivo  directo en el hospedero no debe ser considerados como probi&oacute;ticos y que  deber&iacute;an m&aacute;s bien ser llamados agentes bio-remediadores.</p>     <p>De igual manera, la utilizaci&oacute;n de bacterias Gram-positivas como <i>Bacillus</i>, en general ha demostrado ser ben&eacute;fica en  el cultivo de camar&oacute;n (<i>P.  monodon</i>, <i>L. vannamei</i>), causando un importante incremento en las tasas de supervivencia y peso  (<a href="#tab1">Tabla 1</a>). Sin embargo, pocos estudios han documentado la posible capacidad de  los suplementos probi&oacute;ticos para mejorar la producci&oacute;n de nauplios y poslarvas.  Recientemente Decamp <i>et al</i>. (2008) publicaron  un resumen de los estudios provenientes de Asia y Latinoam&eacute;rica en los que se  utilizaron probi&oacute;ticos para la larvicultura de camar&oacute;n. En ese art&iacute;culo ellos  registraron el desempe&ntilde;o de una mezcla comercial de cepas de <i>Bacillus</i> (SANOLIFE MIC), y encontraron que era una  alternativa pr&aacute;ctica al uso de antibi&oacute;ticos ya que si son implementados de  manera simult&aacute;nea con una buena higiene y medidas sanitarias adecuadas, reducen  el impacto potencial de pat&oacute;genos. Registra tambi&eacute;n que no existen diferencias  significativas entre los tratamientos donde se valoraron algunos aspectos como  la biomasa en el tanque, la supervivencia, la calidad de agua y la carga de <i>Vibrio</i>.</p>     <p>Los principales argumentos para la utilizaci&oacute;n del g&eacute;nero <i>Bacillus</i> como aditivos de piensos para la  acuicultura (Decamp <i>et al</i>., 2006) es que se  trata de bacterias cosmopolitas, que se encuentran en el suelo, el agua dulce y  agua de mar, y tambi&eacute;n en los tractos gastrointestinales de crust&aacute;ceos, peces,  animales terrestres e incluso los seres humanos, adem&aacute;s <i>Bacillus</i> puede producirse en concentraciones muy  elevadas a un costo moderado en comparaci&oacute;n con bacterias que no producen  esporas. Por otra parte, al ser formadores de esporas, son f&aacute;ciles de  transportar y almacenar como liofilizados para ser comercializados. Por otra  parte, las bacterias Gram-negativas son dominantes en el tracto  gastrointestinal de peces y mariscos (Moriarty, 1990; Prieur <i>et al</i>, 1990; Clements, 1997), por lo cual  g&eacute;neros com&uacute;nmente encontrados en estos medios de cultivo como <i>Vibrio</i> y <i>Pseudomonas</i> tambi&eacute;n  han sido evaluadas como posibles probi&oacute;ticos, incluso algunas cianobacterias  han sido propuestas como probi&oacute;ticos para camar&oacute;n (Preetha <i>et al</i>, 2007).</p>     <p>Existen varios productos comerciales, algunos de los cuales son  utilizados en Colombia (MIC Sanolife, INVE; Biostart  Advanced Microbial  Systems, Shakopee, MN; BRF-1A, BRF-13A, PB-32, PBL-44; Enviro-Reps International, Camarillo, CA; PondPro-VC, Biomanagement Systems, Australia; Probiotics, Contessa,ZB Industries, San Pedro, CA; Aquabiotic, Loveland Industries Ltd, USA; Organic GreenTM, Hang Poong Industry, Inchon City, Corea; BioSaf, SafAgri, Minneapolis,MN), principalmente  compuestos por bacterias mirificantes y/o <i>Bacillus spp</i>. las cuales, al disminuir la cantidad  de amonio y nitrito, mejoran el agua, beneficiando la salud de los animales. Es  importante se&ntilde;alar que la calidad del agua de los cultivos intensivos no solo  est&aacute; determinada por la composici&oacute;n de la comunidad microbiana sino tambi&eacute;n por  par&aacute;metros fisicoqu&iacute;micos del agua tales como salinidad, temperatura,  concentraci&oacute;n de ox&iacute;geno, as&iacute; como otros factores estoc&aacute;sticos que pueden  favorecer la entrada y proliferaci&oacute;n de algunos microbios (Verschuere etal, 1997, 2000a; Villamil <i>et al</i>, 2003a).</p>     <p><b>USO DE PROBI&Oacute;TICOS EN EL CULTIVO DE  CAMAR&Oacute;N EN COLOMBIA</b></p>     ]]></body>
<body><![CDATA[<p>En Colombia, a pesar de que la  utilizaci&oacute;n de mezclas bacterianas comerciales en el cultivo de camar&oacute;n es  frecuente en algunas fincas de engorde, hasta el momento existen pocos estudios  cient&iacute;ficos que documenten los efectos reales en el cultivo, de tal manera que  el incremento en la productividad como consecuencia del uso de probi&oacute;ticos en  el pa&iacute;s es controvertido. Existen varias razones que pueden explicar la  discrepancia en los resultados obtenidos ya que hay factores cr&iacute;ticos que  pueden variar los resultados de manera dram&aacute;tica como la dosis del probi&oacute;tico,  el tiempo de administraci&oacute;n, la fase en el ciclo de vida (Villamil <i>et al</i>, 2003a; Mart&iacute;nez-Silva <i>et al</i>, 2008a), adem&aacute;s de la especie seleccionada  como probi&oacute;tico, ya que puede tener actividad antibacteriana frente a un  pat&oacute;geno pero no frente a otro que tenga otras estrategias de patog&eacute;nesis  (Mart&iacute;nez-Silva <i>et al</i>, 2008b).</p>     <p>En los &uacute;ltimos a&ntilde;os, se ha trabajado en  el aislamiento y selecci&oacute;n de bacterias propias del cultivo de camar&oacute;n del  Pacifico, con base en la actividad antibi&oacute;tica in vitro. Los resultados obtenidos con las  bacterias estudiadas muestran resultados positivos durante desaf&iacute;os  experimentales contra bacterias oportunistas como Pseudomonas  aeruginosa (Ram&iacute;rez <i>et al</i>., 2006; Bol&iacute;var, 2008). Es entonces importante continuar con  esta aproximaci&oacute;n metodol&oacute;gica y evaluar la actividad probi&oacute;tica de las cepas  durante infecciones experimentales con agentes infecciosos que ocasionan  importantes mortalidades en el pa&iacute;s, como <i>Spiroplasma sp</i>. (Nunan <i>et al</i>., 2005),  <i>Vibrio spp</i>. (Moriarty,  1998) y virus como la  macha blanca (WSSV) y el Taura, que causan grandes perdidas econ&oacute;micas y  mortalidades (Balc&aacute;zar, 2003). </p>     <p>&nbsp;</p>     <p><b>CONCLUSIONES</b></p>     <p>Los estudios realizados in vivo e in vitro  han demostrado que la  aplicaci&oacute;n de un alto n&uacute;mero de bacterias seleccionadas como probi&oacute;ticos tienen  la capacidad de contribuir al establecimiento de la microbiota intestinal,  incrementar el peso por la mejora en la asimilaci&oacute;n del alimento, incrementar  la supervivencia, la resistencia a infecciones y la respuesta inmune de los  organismos cultivados, as&iacute; como mejorar la calidad de agua en experimentos  realizados a peque&ntilde;a escala. Sin embargo, debido al bajo n&uacute;mero de  publicaciones que describen condiciones de producci&oacute;n intensiva, se evidencia  la gran necesidad de establecer protocolos de aplicaci&oacute;n de mezclas de  probi&oacute;ticos comerciales de tal manera que se garanticen los mejores efectos en  cuanto a supervivencia e incremento en peso en cada uno de los estadios de vida  de los organismos cultivados para entender los episodios de altas mortalidades  causadas por bacterias oportunistas, que han sido registrados en granjas  comerciales. De igual manera, se espera que en Colombia puedan realizarse  aislamientos y selecci&oacute;n de bacterias aut&oacute;ctonas con potencial probi&oacute;tico con  el fin de desarrollar un producto comercial con un protocolo de administraci&oacute;n  estandarizado bajo las condiciones propias de los cultivos del pa&iacute;s, teniendo  en cuanta aspectos tan importantes como la dosis, el tiempo de administraci&oacute;n,  la edad de los organismos cultivados y la escala del cultivo.</p>     <p>&nbsp;</p>     <p><b>BIBLIOGRAF&Iacute;A</b></p>     <!-- ref --><p>1 Abou-Seada, M.y  J. Otto.  1985. Effect of  increasing oxygen concentration on total denitrification and nitrous oxide  release from soil by different bacteria. Biol. Fert. 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