Fermented foods identify cultures and civilizations. History, climate and the particulars of local production of raw materials have urged humanity to exploit various pathways of fermentation to produce a wide variety of traditional edible products which represent adaptations to specific conditions.
Louis Pasteur discussed the issue of fermentation [1,2]. He obviously suspected—and, in some cases, he knew—that fermentative processes are diverse and complicated. Indeed, although modern science recognizes more than one type of fermentation, a general definition should define fermentation as a biochemical process through which most microorganisms decompose carbohydrates to produce energy under anaerobic conditions [3,4,5]. According to Pasteur, “Fermentation is life in the absence of oxygen” [1]. In this somewhat obscure and phenomenological term, various types of fermentation are included, such as yeast fermentation, lactic acid fermentation, butyric acid fermentation, propionic acid fermentation and acetic acid fermentation. The diversity of the phenomenon leads to a variety of end products, which include CO2, ethanol, organic acids and other organic molecules. It is many of these products that make fermentation so useful and interesting both to science and industry [3,4,5,6,7].
The process of carbohydrate decomposition creates a flow of protons and electrons. In aerobic conditions, the final receptor is oxygen (respiration), andin anaerobic conditions, the final receptors are other organic molecules, such as pyruvate or acetyl CoA (fermentation). Fermentation yields much less energy than respiration. The progressive oxidation of sugars involves the transfer of hydrogen ions from intermediate products in the pathway to the final receptor. The Embden–Meyerhof–Parnas pathway (EMP) is utilized in most cases to decompose glucose to pyruvate (homofermentive strains). The Entner–Doudoroff pathway decomposes lactose molecules, while the heterofermentive strains decompose pentoses and other sugars through the phosphoacetylase pathway. The end products depend on the substrate as well as the microorganism species and strain (Figure 1 and Table 1). Different species and different genera have different capacities of fermentation [3,6,7]. For example, E. coli has limited fermentation capacities regarding the variety of end products as compared to lactic acid bacteria (LAB). On the other hand, beneficial fermentation properties can be engineered. By inserting a lactic acid dehydrogenase gene from a bovine muscle, Saccharomyces cerevisiae can produce lactic acid in levels similar tothose of LAB [8].
Table 1. Most common fermentations and the implicated microorganisms
Type of Fermentation | Microorganisms Involved | Food/Environment | Major End Products |
---|---|---|---|
Alcohol fermentation | Yeasts | Wine, beer, sourdough | 2 glucose → 2 glycerol + acetic acid + ethanol |
Lactic acid (homofermentation) | Lactic acid bacteria (Lactobacillus spp. etc.) | Dairy products, fermented meats and fermented vegetables etc. | Glucose → 2 lactic acid + 1ATP (Embden-Meyerhof-Parnas) |
Lactic acid (heterofermentation) | Lactic acid bacteria (Lactobacillus spp. etc.) | 5 and 6 carbon sugars → lactic acid + acetic acid+ ethanol +1ATP | |
Butyric acid | Clostridium spp., Butyrivibrio spp., Bacillus spp. and other anaerobes | Marsh sediments, sewage systems | 4 glucose → 2 acetate + 3 butyrate → butyric acid, butanol, acetoine, isopropanol, acetate, ethanol, 2.3-butadienol |
Mixed acid | Enterobacteriaceae (Escherichia spp., Enterobacter spp., Salmonella spp., Klebsiella spp., Shigella spp. etc.) | Human and animal digestive tract, fresh water | Glucose → acetic acid, formic acid, lactic acid, succinic acid, ethanol |
Propionic acid | Propionebacterium spp., Veilonella spp., Bacteroides spp., some Clostridia spp. | Dairy products | Glucose, glycerol, lactate → propionate, acetate, |
Acetic acid | Acetobacter spp., Gluconobacter spp., Bacillus subtilis |
Acetic acid industry | Oxidation of sugars, sugar alcohols, ethanol →acetic acid |
Microorganisms and carbohydrates are abundant in nature and fermentation occurs in every anaerobic environment. It follows that it is not a question of “if’ but a question of “when” and “how” early humans discovered and harvested the benefits of fermentations in the domain of food and beverage. Near Lake Neuchatel in Switzerland, archaeologists discovered the remains of cheese manufacturing from 6000 BCE [9]. Homer refers to Cyclops Polyphemus as a cheese producer, while in many cultures, such as the Mongols, the Tuareg, the Bedouin, the Tibetans many central Africa tribes and Amazon natives, among others, strong evidence exists of fermented curds production in the antiquity [10,11]. Bread is depicted in ancient Egypt scrolls aging from 2700–2500 BCE. In the third century CE, 72 kinds of bread were listed in Athens [11]. Finally, wine must have been “invented” somewhere between the Black Sea and the Caucasus mountains from 6000 to 3500 BCE. Apparently, someone accidentally left crashed grapes in a container and tasted them much later [12,13,14]. A similar theory is presented for beer brewery, but in this case, meshed cereals with water were left the container [15]. In ancient Egypt, the kneading and baking facilities were beside the beer breweries, obviously both using Saccharomyces cerevisiae [15,16].
One could argue that microbiomes can protect their food ecosystems themselves. Indeed, it seems that microbiomes can deploy certain defence lines and hence rapidly transform their environment in such a way that it finally becomes hostile to most other microorganisms. The key words are “finally” and “most.” It is true that fermentation itself serves as the first line of defence by rapidly decreasing the pH values (the involved microorganisms are also called “fast acidifiers”) through the production of organic acids (e.g., lactic acid) and also by eliminating the carbohydrates of the environment, thus depriving the competitive bacteria of nutrients. However, this process—no matter how fast it proceeds—takes some time, and this provides a window of opportunity to various spoilage intruders or pathogens. This is particularly true for fresh cheeses of artisan origin. Various Brucella strains manage to survive in such products, causing serious foodborne illnesses, although they are eliminated in cheeses with a ripening period longer than 2 months [210]. Normally, by the end of the ripening process, most pathogens and spoilage bacteria will be destroyed or permanently inactivated due to the low pH, deprivation of nutrients and antibacterial activity of certain substances produced by the fermentation flora. Staphylococcal species, on the other hand, survive throughout the fermentation process, though a healthy microbiome can control effectively their populations [211].
Fermentations dominate the history and present of nutrition in the sense that, in one form or another, most humans eat (or drink) fermented foods daily. There is a vast variety of fermented foods and drinks throughout the world, some of which were indicatively discussed. The multitude of microorganisms involved in fermentations—associated with complicated metabolic pathways—is more than impressive. Apart from their nutritional effects, fermented foods enhance and protect the health of the consumer in more than one way. Most of these microorganisms extend their action into digestive tract of the consumer, conferring serious benefits to his/her health. This probiotic effect has been extensively verified in vivo and in vitro by numerous reports. Regular consumption of fermented foods reduces the risk of certain types of cancer, protects the health of the gastrointestinal tract and the urogenital tract, alleviates symptoms of diseases such as Crohn’s disease and irritable bowel disease and enhances the immune system. There are strong indications of a positive impact to certain psychological and behavioural disorders. There is also a certain beneficial effect of their daily use to the oral health. By lowering the pH value in the oral cavity, they prevent dental caries and the erosion of teeth while contributing to the treatment of periodontitis and halitosis. The lowering of the pH value of the fermented products is a defence mechanism of the microbiome, since most antagonistic bacteria cannot survive at low pH values. Bacteriocins are an effective means of overcoming other bacteria. These substances are a very promising tool in the food industry because they are a natural way of preservation, fully compatible with consumers’ expectations for a mild treatment of food. Yet, the increasing incidence of resistant strains in the microbiomes is a trend that should be addressed by the scientific community.
This entry is adapted from the peer-reviewed paper 10.3390/foods10010069