**7. Conclusion and future perspectives**

As stated by Van der Giezen in 2009 "over the last 5–10 years, it has become apparent that the organelle known as the mitochondrion is a much more fluid entity than generally believed," so "why should mitochondrion be the same in all eukaryotes while other cellular structures show such great evolutionary malleability?"

It is our belief that since natural selection has given invertebrates the opportunity to evolve in quick steps, a large window is opening in the field of mitochondrial research among these species, giving an outstanding opportunity to researchers to contribute to an increase in knowledge, not only because there is scarce information, but also because many species have shown special and unique characteristics that need to be explained.

At this point, the information reviewed clearly shows that invertebrates display remarkable physiological capabilities, including highly specialised mechanisms for adjusting mitochondrial functions to solve their energetic demands under the stressful conditions they usually face. These species also include within their systems ancient and novel molecules and structures acting to reach an adaptive state, from the increasing number of mitochondria per cell to the highly complex function of the HIF system.

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It is also remarkable that the number of invertebrate species considered as potential models in the study of mitochondrial function has increased. New data on marine invertebrates, such as molluscs and crustaceans and non-*Drosophila* species, are emerging. Since there is still an immense lack of knowledge about invertebrates, important efforts in new animal models should focus on i) the description of mitochondrial systems in species inhabiting extreme environments, ii) the recognition and understanding of the causes and effects of mitochondrial disorders, and iii) the development of unsolved phylogenetic relationships among species and phyla. This may also open important opportunities for new biotechnological applications to better face the effects of global changes such as warming, hypoxic conditions and chronic stressors that specifically affect the central metabolic pathways in such species.

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*Milano, Italy* 

**Optimisation of Cell Bioenergetics** 

Diego Mora and Stefania Arioli

**in Food-Associated Microorganisms** 

*University of Milan, Dipartimento di Scienze e Tecnologie Alimentari e Microbiologiche,* 

Microorganisms display a considerable versatility, with mechanisms that govern cell bioenergetics and a large number of redox active molecules being used as electron donors or acceptors. We will not review the basis of microbial bioenergetics here, but instead focus attention on the metabolic systems that microorganisms have evolved to optimise the efficiency of cell catabolism and cell energy homeostasis. The mechanisms that act in the regulation of cell bioenergetics belong to the complexity of biological systems in which large networks of metabolic pathways interact to govern the life and responsiveness of cells towards environmental fluctuations. During growth, all microorganisms determine considerable changes in the environmental concentration of nutrients, organic acids and other molecules generated by cell catabolism. As a consequence, microorganisms are constantly faced with different environmental stimuli and stresses. The natural habitats of some microorganisms may fluctuate erratically, whereas others which are more predictable offer the opportunity to prepare in advance for the next environmental change. In this context, microorganisms may have evolved the bioenergetic machinery to anticipate environmental fluctuations by adapting to their temporal order of appearance. Food matrixes represent an example of 'predictable' fluctuating environments, generated by anthropic activities and able to drive the speciation of several microorganisms. The nutrient's richness, and specifically the abundance of mono- and disaccharides that characterise several food matrixes (such as milk and grape juice), have allowed the speciation of lactic acid bacteria (LAB) and yeasts with a high fermentation capacity. The bakers' yeast *Saccharomyces cerevisiae* degrades sugars to two-carbon components – in particular, ethanol – even in the presence of excess oxygen, thus using a fermentation metabolism instead of the energetically favourable respiration metabolism (2 mol versus about 32 mol of ATP per mol of glucose respectively). *S. cerevisiae* alcoholic fermentation has been exploited for several millennia throughout the world in a variety of food processes of crucial importance for humans, such as the making of beer, wine and bread. Moreover, LAB species have partially lost the genetic information need in order to carry out a respiratory metabolism on behalf of a homofermentative pathway in which lactic acid is the primary product, or a heterofermentative pathway in which lactic acid, CO2, acetic acid and/or ethanol are produced (Kandler, 1983). The seemingly simplistic metabolism of LAB has been exploited throughout history for the preservation of foods and beverages in nearly all societies, and dates back to the origins of agriculture. The domestication of LAB strains passed down through various

**1. Introduction** 

