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J. Chem. Tech. Biotechnol. 1982,32,224-232

Biotechnology in Relation to the Food Industry

B. Jarvis and A. W. Holmes

LeatherheadFood RA, Randalls Road, Leatherhead, Surrey KT22 IR Y, UK

Biotechnology has been defined in various ways but is essentially the application of biological systems to the manufacturing industries. By implication therefore food biotechnology is the application of plant, animal and microbial systems to the production and industrial processing of food, through the development of new cultivars and livestock strains, of microorganisms with particular characteristics, and thence of new and alternative food raw materials, additives, processing aids, etc. After the publication in 1980 of the report of a joint working party on biotechnology,l an appraisal was made of the awareness, interest in and potential applications for biotechnology in the food manufacturing industries.2 This appraisal was done, firstly, by questionnaire to food-manufacturing companies, trade associations, etc., second, by follow-up discussions with appropriate academic, research institute and industrial experts in the area of biotechnology and, third, as part of a Delphi forecasting exercise.3 Biotechnology is not new. Indeed, the food and related industries provide potent examples of the traditional application of biotechnology in areas as diverse as the brewing of vinegar and alcoholic beverages, through to the production of cured meats, the application of enzymes to the tenderisation of meats and the production of isomerose from starch. Since foods are biological materials per se, any technological treatments of food are, by definition, applications of bio- technology. There are two primary aspects to food biotechnology. Firstly, the positive application of biological processes in the development of new or improved food pro- ducts and, second, the application of what we refer to elsewhere4 as ‘negative biotechnology’. The latter is the application of technological skills to the prevention of undesirable biological change. Both are relevant to the modern food-manufacturing industry and must be considered in parallel. This paper is concerned specifically with the application of positive biotechnology to the food industry. The objective is to summarize some of the findings from our surveys and to indicate areas in which biotechnology developments may be applied in the food-manufacturing industries.

1. Awareness of food biotechnology

1.1. State of the art in the food industry The industrial food-manufacturing industry can be divided into three clearly defined classes. The first, and numerically the largest, group is a craft-based low-technology sector which shows almost complete unawareness of the existence of modem biotechnology, even in relation to their day-to- day activities. The second group of companies, while not actively engaged in biotechnology, takes a keen interest in developments and would clearly welcome opportunities to use biotechnology developments. It is probable, however, that these companies will be unable to capitalise on oppor- tunities without considerable input of both skilled personnel and capital investment. In product- management terms these companies will comprise the ‘developers’ rather than the ‘innovators’. Finally, there is the small group of high-technology companies within, or associated with, the food industry which already operate successful biotechnology research teams. Many of their develop- ments are well known: for instance, the Rank Hovis McDougall development of a mycological protein (recently approved for human food use); the development of microbial gums by Tate and Lyle; the cloning of palm trees by Unilever Ltd; also the development of enzymes for food applica-

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tion, e.g. Miles Laboratories Ltd and Novo Enzymes Ltd. All of these companies have taken an objective look at the possibilities for application of biotechnology in the food industry. As will be discussed later, they have found that many possibilities fail on purely economic grounds. They have also found major problems of legislation, safety clearance and consumer acceptability in the applica- tion of biotechnology developments in the food industry. These matters will be discussed in more detail later. Generally, the high-technology companies acquire the necessary resources to pursue the possi- bilities which they have identified. Where specialist resources are required, the work may be subcontracted to appropriate university departments or research institutes.

1.2. State of the art in universities and research institutes

By and large the outstanding feature of virtually all university departments visited was the apparent lack of interest in the application of biotechnology to the food industry. In general most university departments are concerned apriori with fundamental studies rather than with potential applications for the systems or products. Within the ARC Research Institutes relatively little biotechnology is related directly to the food- manufacturing industry. Work is being undertaken on: genetic manipulation of dairy starter organ- isms; procedures for isolation of commercially valuable protein fractions from milk and other agricultural commodities; anaerobic fermentation of effluents. However, in the fullness of time it is conceivable that some products developed for clinical application may well find uses in food manufacture.

2. Current food-industry biotechnology processes

Table 1 summarises some of the current areas where biotechnology is being utilised by the food- manufacturing industries. From this it can be seen that, although some of these processes may be considered to be ‘traditional’, many are of relatively recent development. One may therefore con- clude that appropriate parts of the food-manufacturing industry are already heavily engaged in the application of biotechnology. The circulated questionnaire yielded an overall response of 24%. In summary form (Table 2) it may be seen that some 40% of the respondents already actively use enzymes or microorganisms

Table 1. Some current food-industry biotechnology processes

1. Production

(a)

Food additives, such as citric and gluconic acids

(b)

Starch hydrolysates by the use of amylase

(c)

Protein hydrolysates for flavour purposes

(d)

Fermented foods, drinks, etc.

2. Modification

(a)

Sugar confectionery centres by the use of invertase

(b)

Glucose syrups by the use of isomerase to produce high-fructose syrups

(c)

Proteins to increase their functional properties by the use of proteinases

(d)

Meat by tenderisation with papain

(e)

Desugaring of egg white by glucose oxidase

3. Improvement

(a)

Improvement in the technology of cheese production by the use of selected microorganisms or non-natural enzymes

(b)

Use of pectinase to increase the yield and/or clarify fruit juice

(c)

Development of improved flavours by the enzymic modification of dairy fats

4. Other applications

(a)

Detoxification of rapeseed by the hydrolysis of glucosinolates with

endogenous myrosinase

(b)

Analytical use of enzymes, e.g. glucose oxidase

15

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B. Jarvis and A. W. Holmes

Table 2. Industrial responses to the biotechnology questionnaire

Companies which:

Respondents"

(%)

-

A.

Use enzymes or microorganisms in processing

41

B.

Use

food additives/processing aids

82

C.

Can identify potential supply problems which might be alleviated through biotechnology

34

 

(40 % of B)

D.

Would be prepared to utilise microbial proteins if commercially available

24

E.

Are involved in up-grading by-products such as skim milk and whey

24

F.

Are interested in upgrading food wastcs by biotechnology

46

G.

Are interested in on-site fermentation of food wastes to produce methane or gasohol

31

H.

Are already involved in F or G

22

J.

Are involved in, or proposing to do, biotechnology R & D

33

K.

Consider that biotechnology will expand or improve the profitability of their operations

41

Of 680 questionnaires circulated 163 (24%) were returned.

(+ 27 % unsure)

in their processes and that 82 % already utilise various types of food additives and processing aids, some of which may already be derived by biotechnology processes. Of these, approximately half identified potential supply problems of food additives and processing aids, which might well be alleviated through biotechnology developments. Forty-one per cent of the respondents considered that biotechnology would definitely expand or improve the profitability of their operations if applied appropriately; 32% did not believe that biotechnology had anything to offer to their operations; a further 27% were uncertain. While it is tempting to extrapolate these data to the food industry as a whole, it would be unwarranted since it may be supposed that a high return will have been achieved from those companies with an interest in biotechnology. However, it is likely that the actual use of enzymes in food processing is far greater than was recorded in the questionnaire responses. For instance, the Association of Microbial Food Enzyme Producers identifies 29 different enzymes for use in no less than 18 different parts of the food industry, and it should be noted that of these all but four are hydrolytic enzymes. A major potential feedstock for biotechnology is the vast amount of food-industry waste material. Some 73 respondents provided data on the sales value and disposal costs of liquid and solid waste; these are summarised in Table 3. Waste packaging materials and other solid wastes account for a relatively small proportion of the total and much is already sold by the industry. By contrast, liquid effluent is available in considerable quantity and the costs of disposal are high. It might be supposed that such material could readily form feedstock for biotechnology processes but in many cases the pH and the concentration of fermentable solids are such as to make the effluent unsuitable for feedstock use without a major modification of upstream processing to divert and separate heavily contaminated and lightly contaminated process waters. If segregation can be undertaken, or dilute effluent concentrated at low cost, the possibilities for feedstock use must be high, particularly in relation to energy and biomass production.

Table 3. Some food-industry waste materials

 

Weight

Sales

Disposal

 

No.of produced Amount sold value

costs

Type

sites

(Mt)

(Mt)

(fm)

(fm)

Liquid effluent

46

29.6

0.001

0.04

5.1

Solid waste

91

6.7

6.3

145.1

0.4

Packaging material

56

0.02

0.015

0.2

0.05

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3. Potential areas for biotechnology developments

The major areas for future development are summarised in Table 4 but this is by no means exhaustive since many possibilities have been excluded. The respondents in the Delphi study3 considered that over the next 10-20 years microbial enzymes will be used increasingly for continuous food processingand for waste treatment. Microbial fermenta- tion will be used for modification of cellulose, detoxification of food materials and upgrading of industrial waste materials. Microbes and their enzymes will be used increasingly for food analysis, especially for detection and analysis of food-associated carcinogens.

Table 4. Potential areas for food biotechnology developments

(I)

Production of:

Proteins Carbohydrates Fats and oils

F1avours

Manufacturing aids

(2) Detoxification of food-associated toxins

(3) Analytical applications

(4) Upgrading of effluents and waste materials

In addition to the production of specific biotechnology products for use in food manufacture, the

panel considered that biomass would be used extensively for animal feeds; however, the probability for widespread use of biomass in human foods was more limited. The major role for biotechnology in relation to proteins would be the development of proteins with specific functional properties.

will be used to generate ‘starter cultures’ with specific attributes

Genetic manipulation of microbes

for industrial fermentations and for food fermentation use. Concern was expressed at the potential

health risks associated with modem versions of some ethnic food fermentations and suggested a need to develop ‘starters’ which might be used to control reliably the growth of food-associated pathogens. The sections below expand on some of the specific developments which are attractive for one reason or another.

3.1. Carbohydrates

3.1.1. Production of polysaccharides The production of microbial gums has been identified by many people in the food industry as an area in which supply problems may be alleviated by biotechnology developments. It must be recog- nised that this response may have been influenced by the knowledge that Tate and Lyle are able to produce xanthan gum and this may have led other companies to believe that the technology is capable of extension. For reasons discussed later, it should be noted that the gums produced by Tate and Lyle are not being used in the food industry, the present major outlet being in North Sea oil drilling. There is no doubt that many microorganisms are capable of producing polysaccharides and it seems very likely that a suitable screening programme could lead to the development of a full range of thickening agents for use in the food industry. The current selling price of such materials is about €1400 t-l and the market value is of the order of E7 million per annum. This annual market might be sufficient to bear the necessary R & D capital-investment costs but if an expensive safety- evaluation programme was required, it is unlikely that this type of development could be justified. An alternative solution would be that of improving the yield of plant gums by the use of enzyme- extraction processes. It is conceivable that such processes might well change the properties of the gums in such a way that alternative applications might arise. For instance, the natural gum may well occur in plant tissues as a mucoprotein, the link between the polysaccharide and protein being

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disrupted in the traditional extraction process. If, by use of enzyme technology, one could extract the mucoprotein intact, it seems highly probable that the properties of the mucoprotein would be significantly different from those of the products which are used today. Furthermore it might be possible to utilise enzyme technology to modify the properties of gums produced by traditional methods. Such approaches would not require such vigorous safety evaluation and may well be acceptable commercially.

3.1.2. Enzymic modificationof other polysaccharides

There is considerable interest in the possibilities for enzymic modifications of carbohydrates lead- ing to the synthesis of starches and other polysaccharides with improved physical properties. The hydrolytic approach has been investigated extensively already and in our opinion it is unlikely that any significant new developments will occur. However, a biosynthetic approach is at least theoretic- ally possible and warrants investigation.

3.1.3. Sugar esters Sugar esters, presently made by complicated chemical processes, are becoming of increasing import- ance as food materials. It is possible to contemplate a situation whereby the coupling of the sugar to the fatty acids is carried out enzymically. Many sugar esters are known constituents of microbial cells and it may well be possible to prepare them directly with genetically manipulated organisms. It is probable that such products would not be required to undergo stringent safety testing to the same extent as products manufactured by chemical processes. However, since heterogeneous enzymic reaction systems would be involved, much fundamental investigation will be required.

3.2. Proteins

3.2.1. Biomass

It has been suggested that a need exists for biomass production on a commercial scale, both for human food and for animal feeds. It is well known that many companies have explored the develop- ment of biomass for animal-feed purposes and such materials must compete on the economic market with other sources of protein. It is our opinion that the economic viability of such processes is limited unless the protein is produced as a by-product of some other biotechnology, e.g. energy production. The availability of single-cell protein for human food is, of course, very different. At the present time UK approval has been given for only one such product, the mycoprotein manufactured by Rank Hovis McDougall Ltd. To reach this stage of development has required considerable invest- ment by RHM and their collaborators. It may be appropriate to consider the attitude which the food industry might take on the use of microbial proteins. If the material is to be used in developing new food products it must offer some positive advantage to the food technologist; such an advantage may be either price or functionality. It is unlikely that biomass can every be produced at a price so low as to offer a very large financial advantage over, for instance, soya protein and one must there- fore seek some functional advantage to justify its use. Many microbial proteins are effectively non- functional because the extraction and subsequent treatments denature the proteins. Hence at best the materials will act as fillers. The RHM product appears to offer an advantage in being a low-cost texturised material which lacks the off-flavours normally associated with vegetable proteins. It can be argued therefore that functional advantages may be of greater importance. Since the RHM material is likely to be capable of fulfilling any opportunities which exist for meat and meat-like products, there would seem to be little point in duplication of this product at the present time. One must therefore consider what other functional properties of proteins might be desirable. In different situations proteins are expected to be able to foam, to emulsify or to gel. Traditionally egg proteins have been able to fulfil these requirements but increasingly new materials prepared by fractionation of vegetable and milk proteins are becoming available which offer real advantages over egg proteins. Until these techniques have been fully exploited, there seems little point in pursuing further investigations in this area.

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3.2.2. Milk proteins

Many industrial companies expressed concern about the problems of disposing of whey and skim milk and/or the desirability of producing useful materials from these dairy by-products. However, this is an area which is already heavily researched by many organisations both in the UK and abroad and protein fractions are appearing already on the commercial market.

3.2.3 Alternatiue proteins

It has been suggested that if a satisfactory market existed for, e.g., serum albumin it would be possible to produce such a protein from microbial sources in sufficient quantities to service the food industry. However, because of the low yields from microorganisms it seems unlikely that this will ever be a viable proposition for the food manufacturer. Furthermore serum albumin and other serum proteins can be produced by fractionation of unwanted blood available in large quantities at abattoirs. Work on the texturisation of blood proteins is being studied extensively, notably in the Department of Food Science at Nottingham University. Similarly, although it is theoretically possible to produce gelatin from microbes, the present state of the gelatin industry indicates that this is unlikely to be a viable process.

3.2.4. Meat tenderisation

Several companies identified the need for improved tenderisers for meat and one recognised that the requirement was the specific softening of connective tissue. It is possible to contemplate a situation in which elastase is coupled to a carrier with a specific affinity for elastin so that one can achieve specific softening of connective tissue. Such work would require fundamental knowledge of techniques for coupling enzymes to carriers but if successful could be used to upgrade consider- able amounts of meat which at the present time are suitable only for manufacturing purposes.

3.3. Fats and oils

3.3.1. Production offats

There is undoubtedly a need to produce fats and oils with particular characteristics; for instance, to serve as partial replacements for cocoa butter, palm oil, illipe and shea fats. The work of Ratledge5 and others demonstrates that the possibilities for production of fats from plant and microbial cells are high but they must be related to the relative costs. The work of Ratledge suggests that at 1978 prices the costs of producing fat by biotechnology may be high as €700t-1 and it follows therefore that unless the traditional material costs more than ElOOO t-1 there is no point in considering its production by microbial fermentation techniques. Illipe and shea fats cost up to €1500 t-1 and are available only in restricted amounts. Cocoa butter costs upwards of €2000 t-1 at current prices. In these situations therefore it is conceivable that microbial production of fats could be justified on purely economic grounds; but there is no economic justification for producing palm oil by biotechnology since it currently sells at about 400t-l. Although algae are reported to produce up to 40% of their dry weight as fat, we do not feel that suggestions for production of fat by algae are tenable in the UK.

3.3.2. Fat extraction from oilseeds

It was suggested that the use of solvents to extract fat from oilseeds might eventually become unacceptable and that it is desirable to develop an alternative technology based on the use of enzymes. While such a technology is theoretically possible, it is unlikely that it would ever find favour since, in the event of solvent extraction becoming unacceptable, it is likely that a hot crushing procedure would be brought back into operation.

3.3.3. Hydrogenation and dehydrogenation reactions

At the present time the properties of fats and oils are modified by the chemical process of catalytic hydrogenation, in the course of which trans-acids may be produced. One can conceive of a process

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in which the hydrogenation is undertaken by microbial or enzymic processes which may not give rise to the formation of trans-acids. A major problem will be the generation of an appropriate reducing system at reasonable cost.

3.3.4. Triglyceridemodification

Modification of oils and fats to produce specific triglycerides present similar problems to those discussed above in relation to starch modification. In addition, it is almost certain that the reaction will have to take place at an interface which will require a better understanding of factors controlling enzyme reactions in a biphasic system.

3.4. Food flavours

The attraction of producing flavour compounds by biotechnology is great. Two specific flavours which would appear to present supply problems at the present time are vanilla and capsicum and it has been suggested that these could both be prepared by tissue culture of plant cells. In the case of capsicum this would undoubtedly require plant-cell fermentation techniques which are expensive in relation to product yield. Since the flavour is a mixture of materials it is most unlikely that the genetic information could be identified and transferred to a more rapidly replicating cell system such as a microbial system. Even with plant cells it may be difficult to persuade the cells to differen- tiate and produce the appropriate flavours. A similar situation pertains with respect to other flavours of interest; for instance, mint and onion flavours. The prospects for vanilla may well be much more promising since the complexity of the flavour compound is very much lower and it may be possible to transfer the relevant genetic information to an appropriate microorganism. Clearly there are inherent technical difficulties in this area and much more fundamental work is required in the area of multiple genetic transfer, particularly from plant and animal cells. Nevertheless, the rewards in this area may be enormous if the problems could be overcome.

3.5. Food colours

A further group of compounds of considerable interest from a biotechnology point of view is that of colouring materials. There is no doubt that some microorganisms are capable of producing various colourings and one was on display in Kyoto at the 1978 World Food Congress. However, merely because the material is synthesised by microorganisms it cannot be regarded as natural and, although it is possible to produce some nature-identical materials such as anthocyanin by plant culture, the concentration of these materials under appropriate conditions would rise only to about 10% of cell dry weight. This means that material could cost anything up to €300 OOO t-l of 100% pure material and such a price is hardly competitive with a natural product. It is, of course, possible to contemplate the transfer of genetic information from plants to microorganisms and in this situation the economics might well be very different. A further possibility is that of enzymic modifi- cation of anthocyanins and other colour precursors to yield a range of semisynthetic colour pigments. The whole of this area, however, is faced with problems associated with the need for extensive toxicological testing which might well make such processes totally uneconomic.

3.6. Antimicrobial aspects of biotechnology

The possibilities for suppression of growth of pathogenic and spoilage organisms by the deliberate use of appropriate types of desirable organisms, is of considerable interest in many areas. There is a tradition of such an approach in the production of lactic dairy products, fermented meats, etc. Nevertheless, the selection of microbial strains with appropriate characteristics is not easy and, for food applications, will undoubtedly require genetic manipulation of selected organisms to transfer all the desirable characteristics from a group of organisms into a single organism. If it were possible to satisfactorily develop such an organism, it could have a substantial effect on current food-preser- vation techniques. It would be possible to envisage that freshly butchered meat could be inoculated

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with an appropriate culture, packed and distributed to retail outlets without the need for refrigera- tion. It could also conceivably reduce the need for long-term preservation techniques with canning and freezing and also reduce the need for added chemical preservatives. A number of compounds known to chelate trace metals has been investigated in the clinical context.s It is our belief that compounds of this sort may well have potential application in the food area. Production of natural antimicrobials from plant cells is well documented. If an appropriate system for large-scale plant-cell cultivation could be geared towards production of such anti- microbials, it is conceivable that some might have application as preservatives for food and other materials. None the less, as with colours and flavours, there is a distinct problem associated with the need for toxicological testing.

3.7. Detoxification of foodstuffs

It is possible to contemplate the development of detoxification procedures which may be of real value to industry. For instance, the use of microorganisms or of enzymes specifically to remove mycotoxins from contaminated foodstuffs would be invaluable. Similarly, a process for the removal of erusic acid from rapeseed oil and of glycosinolates from rapeseed meal would be useful, though less economicallyjustifiable.

3.8. Analytical applications

The use of enzymes and serological reagents for analytical purposes is well established both in the food and other areas of industry. The scope will undoubtedly increase with availability of a wide range of such reagents. In addition, appropriate enzymes may be of value in degrading foodstuffs in order to release trace metals or vitamins for subsequent analysis. It should be possible also to utilise a composite enzyme treatment to simulate digestive processes and thereby provide a better estimate of the dietary fibre content of foods. Radioimmunoassay techniques also have a much wider potential than has been utilised to date. For instance, they might be applied to the determination of the lean-meat content of meat products and to the detection and quantification of microbial toxins. Enzyme-linked immunoassays may offer advantages of high sensitivity in such types of applications.

3.9. Other areas

Reference has been made already to the upgrading of effluent and industrial waste and much work on small-scale processes is already being undertaken both in the UK and elsewhere. If such upgrad- ing can yield both a saleable end product, e.g. biomass for use as animal feed, together with gasohol as a source of energy for heating purposes, clearly this would have many advantages to industrial operators, not only in the food-manufacturing industry.

4. Constraints affecting biotechnology developments in relation to the food industry

The major constraints can be identified as: (a) economic; (b) technical feasibility; (c) toxicological acceptability; (d) consumer acceptability.

4.1. Economic

From our calculations raw-material production based on microbial biomass is uneconomic unless the natural material cost is greater than E2000 t-1 at 1980prices. Forlow-volume metabolic products produced by plant-cell culture and critical cost is in the order of El00 OOO t-1. Such constraints may be of lesser significance if political or other supply problems exist. These calculations make no allowance for capital investment and R & D costs; in the present economic climate it is difficult to envisage the use of such capital for development of food-oriented biotechnology, except in a very few restricted areas.

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4.2. Technical feasibility

Although many short-term developments are possible, scientific and technological advances are required in many areas to permit capitalisation by the food-manufacturing industry on biotechnology opportunities. Such technical feasibility studies should be undertaken, however, since if attempts to innovate at this time are not made the European industry will rapidly fall behind many other countries in the developed world. In many areas, short-term developments should be possible with- out resort to genetic manipulation; however, a need exists for research into multiple gene transfer with particular reference to production of complex metabolites typical of many food additives. Further work on transfer of genetic information from eukaryotes to prokaryotes is also required.

4.3. Toxicologicalacceptability

At the time our survey was undertaken severe reservations were expressed concerning the cost and time involved in obtaining safety clearance from MAFF and DHSS and we recommended that clear guidelines should be laid down by these Government departments concerning biotechnology developments relevant to the food-manufacturing industry. A step towards satisfying this need has already been taken in relation to the procedures for clearing novel foods. None the less, there is still considerable scepticism in those parts of the industry concerned with novel foods and food ingredients, Concerning the long-term practicability of undertaking the necessary toxicological studies to satisfy the requirements of Government departments.

4.4. Consumer acceptability

Wherever technology leads to the development of novel foods and/or food ingredients including additives, it will be necessary to sell to the consumer the concept that biotechnology products are not harmful and, if possible, that they offer specific benefits to the consumer. Even when a product has been demonstrated to be safe for food and biotechnology plant operatives, the consumer must be convinced also that he/she is not being asked unwittingly to consume materials which may be dangerous, nor to move to a state where the principal item of the diet-will be ‘pills’. It is important therefore that the consumer organisations are kept fully informed as developments occur so that they can critically evaluate and hopefully provide a rational judgment as to the appropriateness of biotechnology developments in the food-manufacturing industry. With an open approach such as this, the suspicions of consumer organisations will be allayed and consumer resistance minimised.

Acknowledgments

This study could not have been completed without the assistance of many people both within the RA and in the various research institutes, universities and industrial companies. We thank all who contributed and, in particular, C. R. Elson, Dr P.A. Gibbs and Dr J. B. Rossell of the Leatherhead Food RA who undertook some of the visits and discussions. The assistance of M. Barnfield and A. G. Crawford in analysing the questionnaire returns is gratefully acknowledged.

References

1. Anon. Biotechnology-Report

2. Holmes, A. W.; JaNiS, B. Application of Biotechnology to the Food Industry-an

of a Joint Working Party HMSO, 1980.

RA, 1981.

Appraisal Leatherhead Food

3. Jarvis, B. Food microbiology into the 21st century-a Delphi forecast in advances and prospects in food micro- biology. Society for Applied Bacteriology Jubilee Symposium Proceedings July 1981.

4. JaNiS, B.; Paulus, K. J. Chem. Technol. Biotechnol. 1982, 32,233.

5. Ratledge, C. &on. Microbiol. 1981, 11, 263.