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1.

Residuos Agroindustriales para la produccin de Pleurotus Ostreatus


Se llama residuos agroindustrial al material o elemento que despus de haber
sido producido, manipulado o usado a nivel agroindustrial no tiene valor para
quien lo posee y por lo general se desecha no adecuadamente generando
contaminacin en el ecosistema (Atlas, R. y Bartha, R. 2006).

Entre los residuos slidos generados en las ciudades, cerca de 40 % son


materiales celulsicos o lignocelulsicos que, en su mayora, no reciben
tratamiento alguno, como los paales desechables usados y el pasto cortado
de los jardines. (DELFN ALCAL & DURN DE BAZA, 2003).

Los sustratos que se usen para realizar el cultivo del hongo Pleurotus ostreatus.
Deben poseer caractersticas como son: buena disponibilidad en cantidad y
continuidad, buen conocimiento de sus caractersticas fsico-quimicas,
localizacin fcil y cercana, facilidad de transporte y manejo, adems de un
precio ventajoso de adquisicin (Flores y Arias, 2006)

2. Caractersticas de la biomasa lignocelulsica.


Las principales fuentes lignocelulsicas estn conformadas por : [18]
Madera: Bosques vrgenes, plantaciones, residuos primarios de bosques,
residuos de procesa-miento secundarios
Residuos agrcolas: De cereales (trigo, arroz, cebada), bagazo (caa de azcar,
sorgo dulce), ras-trojos o ameros (maz), nabo forrajero.
Residuos municipales
Residuos de papel
Micro y macro algas.
La lignocelulosa est presente como un bloque dentro de la estructura de la
pared celular de la planta. Los principales constituyentes de la lignocelulosa
son la celulosa, la hemicelulosa y la lignina. A conti-nuacin se presenta un
esquema con las componentes de la madera:

3. Composicin Qumica de los Residuos Agroindustriales


Los materiales utilizados para el cultivo de Pleurotus ostreatus, estn
constituidos de compuestos ligninocelulosos, los cuales estn formados por
celulosa y hemicelulosas enlazados mediante lignina, un polmero aromtico
altamente oxigenado, con un esqueleto de fenilpropano que se repite. Sobre
esta matriz se deposita una mezcla de compuestos de bajo peso molecular
llamados extractivos. (Quindi, 2013)

La pared celular vegetal est compuesta bsicamente de celulosa,


hemicelulosa y lignina, las cuales les brindan rigidez y proteccin a la
planta.
4. Material Lignocelulosico
All LC materials have structural similarities. Therefore, a model LC (wheat
straw) was chosen as the substrate to test a series of different chemical and
physical treatments. The effect of these treatments was measured by chemical
and biological assays which indicated the value of the treated material for both
saccharification and animal feed purposes. At the end of the study a number of
different LC were used in comparative tests to determine whether results
obtained from one monocotyledonous model substrate (wheat straw) could be
extrapolated to other monocotyledonous LC as well as dicotyledons. Some of
the key facts about wheat straw, the chosen model substrate, are given below.
(CASTRO, 1994)
4.1.Celulosa
Es el compuesto ms simple encontrado en el material lignocelulosa de las
plantas, es el polmero ms abundante en la bisfera. Est compuesto por un
polmero de residuos de D-glucosa unidos por en laces -1,4. Debido a su
estructura las cadenas de celulosa estn unidas por puentes de hidrgeno
intermoleculares que forman agregados (microfibrillas). (Quindi, 2013)
La celulosa es una molcula que da estructura y soporta a la planta y forma un
cristal empaquetado que es impermeable al agua, por lo cual es insoluble en
agua y resistente a la hidrlisis (Atlas, R. y Bartha, R. 2006). Los hongos
Macromycetes pueden degradar la celulosa por medio de la produccin de
enzimas como son endo--1,4-glucanasa, el complejo Cx y endo--1,4glucosidasa (Martin, A. 2005).

La celulosa es un componente fibrilar constituida por un polmero lineal de


restos de D-glucosa unidos por enlaces (1-4) glucosdicos.
Cada molcula presenta una rotacin de 180 grados con respecto a las
molculas contiguas, estabilizadas por la formacin de puentes de hidrgeno
intramoleculares entre OH-3-O-5 y OH-2-OS-6. La ausencia de cadenas

laterales unidas a la estructura lineal de las cadenas de (1-4) glucano,


permite la formacin de agregados moleculares (microfibrillas) estabilizados
por puentes de hidrgeno intermoleculares entre 0H6-0S3, confirindole una
estructura cristalina. (Fernandez,F. 2004)

La celulosa es un componente mayoritario de la madera y otras fibras


vegetales, cadenas polimricas que al ser hidrolizadas liberan unidades de
glucosa. Asociadas con las fibras celulsicas hay hemicelulosas, que son
polmeros de azcares distintos a la glucosa, bsicamente xilosas y manosas
(Eyzaguirre 2000). La hidrlisis de celulosa y hemicelulosa depende de la
estructura de la que forman parte, de los monmeros que las conforman y del
tipo de enlace entre ellos (Hadar et al. 1992).

Cellulose is the most abundant cell wall component in plants. The basic
structure consists of a linear polymer of up to 14,000 anhydro glucopyranoside
units linked by -1,4-glycosidic bonds (Cowling and Kirk, 1976). These linear
polymers in turn aggregate to form fibrils. The structure of cellulose will be
partly dependent on the source of the cellulose. For example, during initial
stages of primary cell wall growth cellulose fibrils (primary cellulose) with
approximately 60-70 glucan chains are formed (Preston, 1974). Secondary cell
wall fibrils are considerably thicker, i.e.
aggregation of primary fibrils. The degree of polymerization can be also vary
between primary and
secondary celluloses (<4,500 and 14,000, respectively) (Blaschek et al., 1982;
Marx-Figini,
1966). These long non-branched chains are deposited in an antiparallel
arrangement and bound
laterally through numerous hydrogen bonds (Dey and Brinson, 1984). The
stereochemistry of the
glucose units in cellulose chains, with each glucose moiety being translated
180o to its neighbour,
permits three hydrogen bonds per residue between each adjacent chain (Fig
1.3.1.1).

This degree of hydrogen bonding is unique in carbohydrate structures and is


the prime
reason for the strength of the cellulose molecules. The extent of association
between individual
chains will confer their degree of parallelism or crystallinity index (Cowling and
Kirk, 1976). Both crystalline (highly oriented) and amorphous cellulose are
present in plants, the former structure being more commonly observed.
This highly ordered three-dimensional structure confers the mechanical
strength of cellulose, see above, and also results in its low susceptibility to
chemical and enzymic attack. The disruption of this structure requires severe
treatment conditions which can result in a substantial improvement of cellulose
accessibility to chaotropic agents. However, the intrinsic characteristics of
cellulose alone are not the only factor limiting its accessibility. Existence of
external components, e.g. lignin-hemicellulose structures which are closely
associated with cellulose, can also play an important role in determining the
extent of cellulose utilization by, for example, enzymes and microorganisms
(Cowling, 1975).
Physical treatments, e.g. ball milling, which affect native characteristics of
cellulose (e.g. degree of polymerization and crystallinity index) improve its bioutilization but limiting chemical
barriers (i.e. lignin bonding) remain. Chemical treatments, e.g. alkali, acidic and
oxidative processes, can disrupt both the native cellulose structure and the
associated steric lignin barriers resulting in enhanced bio-utilization (Millett et
al., 1975).
4.2.Hemicelulosa

La hemicelulosa est formada por cadenas cortas y son polmeros


heterogneos que contienen tanto hexosa (azcares de 6 carbonos como
glucosa, manosa y galactosa), como pentosas (azcares de 5 carbonos como
xilosa y arabinosa). Dependiendo de la especie de la planta estos azucares se
asocian con cidos urnicos formando estructuras polimricas diversas que
pueden estar relacionadas con la celulosa y la lignina. Los tres polmeros
principales son los xilanos, mananos y arabinogalactanos (Atlas, R. y Bartha, R.
2006). Los hongos Macromycetes tienen la capacidad de degradar la
hemicelulosa por medio de la produccin de enzimas como son: xilanasas,
galactanasas, manasas, arabinasas y glucanasas (Martin, A. 2005).

Despus de la celulosa, el componente presente en mayor proporcin, es la


hemicelulosa, la cual est compuesta de polisacridos neutros que presentan
una cadena lineal relativamente larga con ramificaciones cortas de varias
clases de azcares como:
Pentosas: xilosas y arabinosas,
Hexosas: manosas, glucosas y galactosas y,
cidos urnicos como: el cido galacturnico y el cido glucurnico. (Leveau,
J. 2000)

La composicin de la hemicelulosa vara dependiendo de la especie y se ha


determinado que pueden unirse mediante puentes de hidrgeno a las
cadenas de (1-4) glucano de la celulosa, formando una red que limita la
separacin de las microfibrillas y proporciona mayor rigidez a la pared.
(Fernandez,F. 2004)

The term hemicellulose was first used to describe any plant polysaccharide
that can be extracted by mild alkaline solutions (Schulze, 1891 cited by
Wilkie, 1979). Although similar in alkali extraction there are many different
types of hemicelluloses, e.g. xylans, mannans, glucans, galactans and
galacturonans, in plants (Wilkie, 1979). Xylans and mannans are the two
most important groups of hemicelluloses present in LC. In general
hemicelluloses have a low degree of polymerization, <200, (Cowling and
Kirk, 1976).
The xylans are the most common hemicellulose and the main non-cellulose
polysaccharides of angiosperms, i.e. monocotyledons and hardwoods.
Xylans are made up of backbones of anhydro xylopyranosyl units linked by
-1-4 glucosidic bonds to which various residues can be substituted.

Possible configuration of the two most common types of xylans, namely


glucuronoarabinoxylan and glucuronoxylan are illustrated in Fig 1.3.2.1.

Glucuronoarabinoxylans are frequently observed in monocotyledons. These


xylans contain both L-arabinofuranosyl units linked by .(1-3) glycosidic
bonds and glucuronic acid, normally present as the 4-O-methyl ether, linked
by .(1-2) glycosidic bonds to the main xylose chain. The third most common
substituent in xylans, normally attached to the C2 and/or C3 of the xylose
chain, is the acetyl group (Thompson, 1983). It appears that the higher the
acetyl substitution the greater the xylan solubility and lower its the ability
to bind to cellulose (Wallace, 1989). The relative concentration of acetyl
groups in hemicelluloses is also important in the context of steam
treatmentof LC as the efficiency of such treatment relies partly on the
release of these acidic groups to act as catalysts for further chemical
reactions [Aronovsky and Gortner (1930) cited by Muzzy et al.
(1983)].
The second most abundant form of xylan are the glucuronoxylans which are
commonly found in hardwoods and have similar structure to that of
glucuronoarabinoxylans, except that arabinofuranosyl units are absent
(Thompson, 1983).

Although considerable differences in sugar composition and degree of


substitution can be observed amongst various xylans present in
monocotyledons and hardwoods, there are still similarities with respect to
their physico-chemical properties and response to treatment. In general
hemicellulose, in particular xylans, can be efficiently solubilized by means
of mild acid-hydrolysis (Wilkie, 1979). To be able to solubilize glucose from
cellulose to a similar extent there is a need to use considerably harsher
acid-hydrolysis or, alternatively, a combination of acid-hydrolysis with
enzymic treatment. The relative ease of extraction of hemicellulosic sugars
by chemical methods has led scientists to consider hemicelluloses as a
possible source of fuel, feed and chemicals (Thompson, 1983; Overend and
Chornet, 1987).
Mannans are the major hemicellulose component in softwoods. Typically
they are harder to extract from the cell wall matrix than xylans.
Glucomannan, the most important mannan present in softwoods, is made
up of a backbone of glucopyranosyl and mannopyranosyl units linked each
other by -1-4 glucosidic bonds (Fig 1.3.2.1). Acetyl groups can be found
linked to C2 and/or C3 of glucose and/or mannose units depending on the
plant tissue, species, etc. As mentioned for xylans, the existence of such
acetyl groups in mannans has important implications on their physical
properties (Wayman and Parekh, 1990).
4.3.Lignina
Es un polmero complejo tridimensional, globular, insoluble y de alto peso
molecular, formado por unidades de fenilpropano cuyos enlaces son
relativamente fciles de hidrolizar por va qumica o enzimtica. Esta molcula
tiene diferentes tipos de uniones entre los anillos de fenilpropano. (Atlas, R. y
Bartha, R. 2006).
La lignina es la responsable de la rigidez de las plantas y de sus mecanismos
de resistencia al estrs y a ataques microbianos. En las plantas la lignina se
encuentra qumicamente unida a la hemicelulosa y rodean las fibras
compuestas por celulosa (Atlas, R. y Bartha, R. 2006). Los hongos
Macromycetes pueden degradar la lignina por medio de la produccin de
enzimas como son lacasa, lignina peroxidasa y manganeso peroxidasa (Martin,
A. 2005).

La lignina es un polmero constituido por restos fenilpropanoides derivados casi


exclusivamente de los cidos p-cumrico, coniferlico y sinaplico, unidos entre
s por enlaces ter (C-O-C) o C-C. La composicin monomrica, as como el tipo
de enlace entre ellos y su organizacin en la macromolcula vara entre las
diferentes especies.

Su presencia en las paredes celulares secundarias, distribuida en los espacios


entre la celulosa y la hemicelulosa, desplaza el agua por su carcter
hidrofbico, aumentando tanto, la resistencia qumica, como la fsica y la
rigidez de las mismas. (Chihara, G. 1993) (Fernandez,F. 2004).

En los vegetales, adems de celulosa y hemicelulosa est presente la lignina,


que es el segundo compuesto regenerable ms abundante en la Tierra. Se trata
de un compuesto que no slo es recalcitrante a la degradacin, sino que
adems, su estructura reticular tridimensional deanillos aromticos enlazados
por tomos de oxgeno, obstaculiza el acceso de las enzimas hidrolticas hacia
las fibrillas celulsicas. La biotransformacin de la lignina esun proceso clave
en el geociclo del carbono, proceso metablico realizado por mecanismos
oxidantes extracelulares. Los organismos ms eficientes como degradadores
de la lignina son los hongos de la pudricin blanca (Higuchi 1990).

Lignin is the main non-carbohydrate component present in the mature plant


cell wall. It is an amorphous, high molecular weight condensed polymer of
phenylpropane units linked by carboncarbon (C-C) and ether (C-O-C) bonds (Fig
1.3.3.1). Lignins are thought to have the two main functions providing; (1)
resistance to microbial attack and (2) mechanical strength to the cell Wall
(Theander and man, 1984).
Lignins are classified according to the distribution of p-hydroxyphenyl, guaiacyl
and syringyl moieties. These are the corresponding structures to the following
lignin precursors: pcoumaryl, coniferyl and sinapyl alcohols (Fig 1.3.3.2).
Guaiacyl lignins, rich in guaiacyl residues, are found in gymnosperms
(softwoods). Hardwood lignins contain both guaiacyl and syringyl groups in a
ratio that can vary from 4:1 to 1:2. The last and probably most complex group
of lignins is that of grass lignins, as it contains all three aromatic residues in
significant amounts (Nimz et al., 1981). Grass lignins are significantly different
to wood lignins in that they contain phenolic acids (PAC), p-coumaric and ferulic
acids and amino acids closely associated with the matrix 'core' lignin (Van
Soest, 1982). The PAC being linked to the lignin by either ether or ester bonds
(Billa et, al., 1993) and to hemicellulose mostly by alkali-labile bonds (Hartley
and Jones, 1978).

The strong covalent bonds in the complex structure of lignin, i.e. C-C and ether
bonds, makes lignin highly resistant to hydrolytic action of acid and alkali (Van
Soest, 1982). These linkages can, however, be disrupted by mild oxidation, e.g.
nitrobenzene oxidation, which is very often used for analytical measurements
(Wayman and Chua, 1979a). The phenylpropanoid alcoholspresent in 'core'
lignin will produce p-hydroxybenzaldehyde, vanillin and syringaldehyde when
subjected to such mild oxidation. Oxidation of p-coumaric and ferulic acids can
also give rise to phydroxybenzaldehyde and vanillin, respectively (Fig 1.3.3.3).

When grasses are subjected to nitrobenzene oxidation, it is observed that most


of the phydroxybenzaldehyde and up to 25% of the vanillin originates from pcoumaric and ferulic acids respectively (Van Soest, 1982). The occurrence of
PAC in grass lignins has important consequences on the efficiency of chemical
treatments designed to improve the bio-utilization of these materials. The ester
bonded PAC in grass lignins can be solubilized when subjected to cold alkali
treatments. Other lignins which do not contain alkali-labile bonds are
unaffected by cold alkali. In order to depolymerize the lignin 'core' of any LC
the chemical bonds, e.g. C-C and ether bonds between lignin precursors need
to be disrupted by, for example, oxidative treatments (Gould, 1984) or high
temperature chemical treatment (Chua and Wayman, 1979b).

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