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International Journal of Food Microbiology Lifestyles of Sourdough Lactobacilli – Do They Matter for Microbial Ecology And

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International Journal of Food Microbiology Lifestyles of Sourdough Lactobacilli – Do They Matter for Microbial Ecology And International Journal of Food Microbiology 302 (2019) 15–23 Contents lists available at ScienceDirect International Journal of Food Microbiology journal homepage: www.elsevier.com/locate/ijfoodmicro Lifestyles of sourdough lactobacilli – Do they matter for microbial ecology and bread quality? T ⁎ Michael G. Gänzlea,b, , Jinshui Zhengc a University of Alberta, Dept. of Agricultural, Food and Nutrional Science, Edmonton, Canada b Hubei University of Technology, College of Bioengineering and Food Science, Wuhan, China c State Key Laboratory of Agricultural Microbiology, Huazhong Agricultural University, Wuhan, China ARTICLE INFO ABSTRACT Keywords: Sourdough is used in production of (steamed) bread as leavening agent (type I sourdoughs) or as baking im- Lactobacillus prover to enhance flavour, texture, and shelf life of bread (type II sourdoughs). The long-term propagation of Sourdough sourdoughs eliminates dispersal limitation and consistently leads to sourdough microbiota that are composed of Microbiota host adapted lactobacilli. In contrast, community assembly in spontaneous cereal fermentations is limited by Evolution dispersal and nomadic or environmental lactic acid bacteria are the first colonizers of these sourdoughs. Bread leavening Propagation of sourdoughs for use as sole leavening agent (type I sourdoughs) dictates fermentation condi- tions that select for rapid growth. Type I wheat- and rye sourdoughs are consistently populated by insect-adapted lactobacilli, particularly Lactobacillus sanfranciscensis, which is characterized by a small genome size and a re- stricted metabolic potential. The diverse fermentation conditions employed in industrial or artisanal Type II sourdough fermentation processes also result in a more diverse microbiota. Nevertheless, type II sourdoughs are typically populated by vertebrate host adapted lactobacilli of the L. delbrueckii and L. reuteri groups. Metabolic traits of host-adapted lactobacilli that enhance competitiveness in intestinal ecosystems also provide technolo- gical functionality in bread making. Examples include formation of exopolysaccharides, arginine-, glutamine- and glutamate based mechanisms of acid resistance, and glycosyl hydrolases that reduce FODMAP levels in sourdough and sourdough bread. In conclusion, consideration of the lifestyle of sourdough lactobacilli facilitates the selection of competitive and functional sourdough starter cultures. 1. Introduction used in most European baked goods with the primary aim to improve bread quality (Brandt, 2007; Pontonio et al., 2017). Sourdough has been traditionally used as leavening agent for bread Different technological aims of sourdough fermentation – leavening, and steamed bread. CO2 production in sourdough is mediated by yeasts acidification, or dough improver – necessitate different fermentation and heterofermentative lactobacilli, and is coupled to acidification conditions. The traditional use of sourdough as sole leavening agent (Brandt et al., 2004). Microbial metabolism in conjunction with the dictates frequent refreshments of the sourdough, or, in microbiological activity of flour enzymes also improves flavour, texture, storage life, terms, fermentation conditions that maintain sourdough microbiota in and nutritional properties of bread (Gänzle, 2014; Gobbetti et al., the exponential phase of growth. Maintaining sourdough microbiota in 2018). The industrial production of baker's yeast allowed its use as a metabolically active state ensures sufficient CO2 production and lea- leavening agent after the late 19th century; by the middle of the 20th vening power (Brandt et al., 2004). Two examples of fermentation century, baker's yeast had replaced sourdough as leavening agent in processes to achieve leavening are shown in Fig. 1. Details of the fer- most applications except in some small scale artisanal bakeries, for mentation process, i.e. the number of stages per fermentation cycle, and specialty products, and in rye baking, where acidification is necessary the conditions with regards to temperature, time, water addition, and to improve baking properties (Capelle et al., 2013). Recognition of the level of inoculum, are poorly documented in the scientific literature and quality of sourdough bread and the efforts of the baking industry to vary widely between different bakeries; however, type I fermentation offer “clean label” products led to a resurgence of sourdough use since processes generally follow the principle that fermentation microbiota the late 20th century. Currently, sourdough or sourdough products are are continuously maintained in a metabolically active state by frequent ⁎ Corresponding author at: University of Alberta, Dept. of Agricultural, Food and Nutritional Science, 4-10 Ag/For Centre, Edmonton, AB T6G 2P5, Canada. E-mail address: [email protected] (M.G. Gänzle). https://doi.org/10.1016/j.ijfoodmicro.2018.08.019 Received 21 June 2018; Received in revised form 9 August 2018; Accepted 18 August 2018 Available online 20 August 2018 0168-1605/ © 2018 Elsevier B.V. All rights reserved. M.G. Gänzle, J. Zheng International Journal of Food Microbiology 302 (2019) 15–23 Fig. 1. Two examples of sourdough fermentation processes used to achieve leavening of bread without addition of baker's yeast. Panel A, three stage sourdough process (Tang et al., 2017). Tang et al. performed fermentations at 30 °C; in bakery practice, the fermentation stages are performed at 22–28 °C; Panel B; sourdough process for production of Colomba (Raimundi et al., 2017). Other type I sourdough fermentation processes follow a different regimen with respect to incubation time and temperature, and inoculum level, however, all sourdoughs that are fermented to achieve leavening of dough without baker's yeast continuously maintain sourdough microbiota in a metabolically active state. refreshments. This also prevents exposure of sourdough microbiota to and Ripari, 2016). Back-slopping of sourdoughs over long time periods low pH (Böcker et al., 1995; Brandt et al., 2004). The use of sourdough provides opportunity for contaminants from even unlikely sources to as sole leavening agent is generally restricted to artisanal or medium establish themselves as stable members of sourdough microbiota. scale bakeries. Industrial sourdough fermentations employ comparable Böcker et al. (1995) proposed that the technological aim of sourdough fermentation conditions to support leavening in combination with ad- fermentation – leavening or acidification – determines the fermentation dition of baker's yeast. Some of the sourdoughs for use as leavening conditions and the composition of sourdough microbiota. The concept, agent have been maintained over long periods of time; documented initially based on characterization of only a few German sourdoughs, propagation of sourdoughs exceeds 100 years. was further developed in several comprehensive reviews (De Vuyst and Fermentation of sourdoughs with high levels of acidity requires Neysens, 2005; De Vuyst et al., 2014; Gobbetti et al., 2016; Vogel et al., different fermentation conditions; these sourdoughs are fermented at a 1999)todifferentiate between type I sourdoughs, sourdoughs used for high temperature for extended periods of time to achieve high levels of leavening; type II sourdoughs, sourdoughs used for acidification; and total titrable acidity (Böcker et al., 1995; Müller et al., 2001; Rosenquist type 0 sourdoughs, spontaneous sourdoughs. Past reviews focused on and Hansen, 2000; Vera et al., 2012; Viiard et al., 2013 and 2016). microbiological aspects and the impact of fermentation conditions on Dough acidification is particularly relevant in rye baking to inhibit rye composition of sourdough microbiota (De Vuyst and Neysens, 2005; De amylases (Brandt, 2007) and it is used predominantly in countries Vuyst et al., 2014; Gobbetti et al., 2016; Vogel et al., 1999) while ne- where rye bread is popular. These sourdoughs do not allow leavening glecting the link between technological aim and fermentation micro- without addition of baker's yeast. Comparable to the use of sourdough biota that was initially proposed by Böcker et al. (1995). as leavening agent, sourdoughs that are fermented for dough acid- The present communication will initially revisit the question which ification and improved bread quality are maintained by continuous lactic acid bacteria populate sourdoughs. Literature data on sourdough propagation to achieve a consistent composition and activity of sour- microbiota covers more than 300 sourdoughs with documented com- dough microbiota (Brandt, 2007; Zheng et al., 2015b). position and use in artisanal or industrial bakeries, allowing to probe In addition to those sourdoughs that are maintained by back-slop- relationships between the technological aim of fermentation and the ping, spontaneous sourdough fermentations are also used in bakeries. composition of sourdough microbiota. In addition, we address the The most common practice is the use of sponge dough; dough that is question where to lactic acid bacteria in sourdoughs come from; i.e. we fermented for more than 4 h after addition of baker's yeast. Sponge will explore whether the competitiveness of lactic acid bacteria in doughs are commonly used in wheat baking and in production of soda sourdoughs relates to their phylogenetic position or their natural ha- crackers (Sugihara, 1978). Spontaneous fermentations of cereals are bitat. Duar et al. (2017b) proposed that lactobacilli have evolved to also used in the production of cereal beverages, porridges, and
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