Studies on the Uptake and Metabolism of Wort Sugars During
Brewing Fermentations 
	MBAA TQ vol. 43, no. 4, 2006, pp.
            265-269   |  
VIEW  ARTICLE
         G. G. Stewart. The International Centre for Brewing
         and Distilling (ICBD), Heriot-Watt University, Riccarton, Edinburgh
         EH14 4AS, United Kingdom.
      Abstract
      Brewers� wort contains five fermentable sugars: glucose, fructose,
      sucrose, maltose, and maltotriose. In most worts, maltose is the highest
      concentration, followed by maltotriose and glucose. These sugars are taken
      up by yeast in a distinct order (or priority), with glucose repressing the
      uptake of maltose and maltotriose. Glucose is taken up without the
      expenditure of yeast metabolic energy, whereas maltose and maltotriose
      uptake requires energy (active transport). The concentration of each
      sugar, together with their relative proportions to one another, influence
      overall wort fermentation rate and extent. There are also differences
      between ale and lager yeast strains with respect to wort sugar uptake
      characteristics. For example, ale strains are less able to utilize wort
      maltotriose than lager strains. In addition, the concentration of wort
      glucose and maltose influence beer flavor. In particular, wort with
      elevated levels of glucose produces beers with high concentrations of
      esters (particularly ethyl acetate and isoamyl acetate). Worts containing
      high levels of maltose, however, produce beers with much reduced
      concentrations of these esters. This is particularly the case with
      high-gravity worts (>16�Plato).
Keywords: esters, glucose, high-gravity wort, maltose, maltotriose,
      osmotic pressure, yeast viability, yeast vitality 
      
S�ntesis
      El mosto cervecero contiene cinco az�cares fermentables: glucosa,
      fructosa, sacarosa, maltosa, y maltotriosa. En la mayor�a de los mostos,
      maltosa tiene la mayor concentraci�n, seguido por maltotriosa y glucosa.
      Estos az�cares son transformados por la levadura en una orden (o
      priori�dad) muy espec�fico, con glucosa represando el consumo de maltosa y
      maltotriosa. La glucosa es absorbida sin gastar energ�a metab�lica de la
      levadura, mientras que la absorci�n de maltosa y maltotriosa requiere
      energ�a (transporte activo). La concentraci�n de cada az�car, junto con la
      proporci�n relativa de uno con los otros, afecta la velocidad de la
      fermentaci�n y la cantidad total fermentada. Tambi�n existen diferencias
      entre especies de levadura de alta y baja fermentaci�n con respecto a su
      absorci�n de diferentes az�cares. Por ejemplo, levaduras de alta
      fermentaci�n son menos capaces de utilizar maltotriosa del mosto que
      levaduras de baja fermentaci�n. Vale notar que la concentraci�n de glucosa
      y maltosa en el mosto influye sobre el sabor de la cerveza final; mostos
      con altos niveles de glucosa producen cervezas con altas concentraciones
      de esteres (especialmente acetato de etilo y acetato isoam�lico). Mostos
      que contienen altos niveles de maltosa producen cervezas con una mucha
      menor concentraci�n de estos esteres. Esto es particularmente verdad para
      el caso de mostos �high-gravity� (>16�Plato).
Palabras claves: 
      esteres, glucosa, mosto �high-gravity�, maltosa, maltotriosa, presi�n
      osm�tica, viabilidad de levadura, vitalidad de levadura