Section V

The family Leuconostocaceae

Akihito Endo1 , Leon M.T. Dicks2 , Johanna Bjrkroth3 and Wilhelm H. Holzapfel4
1
Functional Foods Forum, University of Turku, Turku, Finland
2
Department of Microbiology, University of Stellenbosch, 7600 Stellenbosch, South Africa
3
Department of Food Hygiene and Environmental Health, University of Helsinki, Helsinki, Finland
4
School of Life Sciences, Handong Global University, Pohang, South Korea, Insheimer Strasse 27, D-76865 Rohrbach,
Germany

The family Leuconostocaceae consists of four genera at present, Leuconostoc (type genus), Fructobacillus, Oenococcus
and Weissella. The phylogenetically closest family is the Lactobacillaceae. Because of a long branch in the 16S rRNA
phylogenetic tree (Figure V.1), Oenococcus species were regarded as rapidly evolving organisms (Yang & Woese, 1989).
This hypothesis was supported by comparative genome analyses of different species of lactic acid bacteria (LAB),
including Oenococcus oeni (Makarova et al., 2006). The family generally forms a robust cluster even if sequences of
housekeeping genes are used for phylogenetic analysis (Figure V.2).
Morphologically, rod-shaped species and coccoid species are present, and the genus Weissella especially includes
both morphologies (Collins et al., 1993). In addition, it shows the widest diversity of peptidoglycan types (Table V.1).
Biochemically, some species are asaccharolytic and sometimes need specific growth factor(s), for example tomato
juice factor for O. oeni (Garvie & Mabbitt, 1967). The family members are obligately heterofermentative LAB and
usually produce lactic acid, carbon dioxide and ethanol during glucose metabolism by the pentose phosphate pathway.
However, the family contains fructophilic LAB, for example Fructobacillus species, which produce acetic acid instead
of ethanol (Endo & Okada, 2008). Fructobacillus species need an electron acceptor for glucose metabolism such as
pyruvate, oxygen or fructose (Endo et al., 2011). The members of Leuconostocaceae are non-thermophilic, and optimal
growth temperature is 2030 C. They are usually non-acidophiles, but O. oeni is a representative of acidophilic LAB,
with strains showing an ability to grow even at pH 3.7 (Garvie, 1967). For growth, the members of the family usually
prefer anaerobic conditions, in contrast to Fructobacillus species, which show a preference for aerobic conditions much
more than anaerobic conditions (Endo et al., 2011). Some of these features can be used as differential characteristics
among the genera of the Leuconostocaceae and obligately heterofermentative lactobacilli (see Table V.1).
The habitats are various. The organisms are usually found in various plants and related fermented products, meat
products and sometimes in the gastrointestinal tracts of animals. Some species are well known as beneficial organisms,
especially for food fermentation and in applications for biopreservation and as desirable flavour producers (Lonvaud-
Funel, 1999; Hornbaek et al., 2004). Because of Psychrotrophic (or psychrotolerant) characteristics of some species,
some strains of these species may cause spoilage in chilled food (Vihavainen & Bjrkroth, 2009). Another negative
aspect is that in clinical practice human infections by vancomycin-resistant Leuconostoc species have been reported
(Janow et al., 2009). However, in no cases have direct relationships between Leuconostoc species and human diseases
yet been proven (Ogier et al., 2008).
In this section, the phylogenetic relationships, phenotypic diversity and practical importance of the members of the
family Leuconostocaceae are described.

Corresponding author email: akihito.endo@utu.fi

Lactic Acid Bacteria: Biodiversity and Taxonomy, First Edition. Edited by Wilhelm H. Holzapfel and Brian J.B. Wood.
2014 John Wiley & Sons, Ltd. Published 2014 by John Wiley & Sons, Ltd.
378 THE FAMILY LEUCONOSTOCACEAE

Leuc. inhae (AF439560)

Leuc. gelidum (AB022921)
Leuc. gasicomitatum (AF231131)
Leuc. carnosum (AB022925)
Leuc. kimchii (AF173986)
Leuc. holzapfelii (AM600682)
Leuc. citreum (AF111948)
Leuc. lactis (AB023968)
Leuc. palmae (AM940225)
Leuc. mesenteroides (AB023242)
Leuc. pseudomesenteroides (AB023237)
Fru. tropaeoli (AB542054)
Fru. pseudoficulneus (AY169967)
Fru. ficulneus (AF360736)
Fru. durionis (AJ780981)
Fru. fructosus (AF360737)
Leuc. fallax (AF360738)
O. oeni (AB022924)
O. kitaharae (AB221475)
W. paramesenteroides (AB023238)
W. thailandensis (AB023838)
W. hellenica (AB023240)
W. confusa (AB023241)
W. cibaria (AJ295989)
W. viridescens (AB023236)
W. minor (TAB022920)
W. halotolerans (AB022926)
W. koreensis (AY035891)
W. kandleri (AB022922)
W. soli (AY028260)
W. ghanensis (AM882997)
Lc. lactis (AB100803)
S. pyogenes (AB023575)
Pl. selangorensis (AF049745)
Ped. acidilactici (AJ305320)
Lb. plantarum (X52653)
Lb. sakei (AM113784)
Lb. casei (D86517)
Lb. reuteri (X76328)
Lb. delbrueckii (AY050172)
Lb. acidophilus (M58802)
Bacillus subtilis (X60646)
0.1

Figure V.1 Phylogenetic relationships among the family Leuconostocaceae and related taxa based on 16S rRNA gene sequences. The tree
was constructed by the neighbour-joining method. Bacillus subtilis was used as an outgroup. Bootstrap values are given at branching points;
values above 70% are indicated

Table V.1 Differential characteristics among the genera in the Leuconostocaceae and heterofermentative lactobacilli (Lactobacillus
group III)

Leuconostoc Fructobacillus Oenococcus Weissella Lactobacillus group III

Cell morphology Cocci Rods Cocci Cocci or rods Rodsb to coccoid

Lactic acid conguration D D D D or DL DL
NH3 from arginine + or + or
Dextran productiona + or + or + or
Ethanol from glucose + + + +c
Effect of oxygen on growth Signicant in Enhanced Suppressed Not signicant Suppressed or not
species possessing signicantc
haem-dependent
respiration
Mol % G+C content in the DNA 3743 4344 3743 3745 3555
Peptidoglycan type Lys-Ser-Ala2 , or Lys-Ala Lys-Ser2 or Lys-Ala, Lys-Ala2 , L-Lys-D-Asp,
Lys-Ala2 Lys-Ala-Ser Lys-Ala-Ser, L-Orn-D-Asp, m-Dpm
Lys-Ser-Ala2 ,
L-Lys-Ala(L-Ser)-L-Ala,
Lys-Ala-Gly-Ala2
a Extracellularly
synthesized homopolysaccharides (e.g. dextran) are produced by heterofermentative lactobacilli associated with cereal environments (see
Chapter 19 on the genus Lactobacillus).
b Except Lactobacillus equigenerosi and Lb. gastricus; they have coccoid cells.
c Except Lactobacillus kunkeei. The species has characteristics similar to Fructobacillus spp.

m-Dpm, meso-diaminopimelic acid.

 THE FAMILY LEUCONOSTOCACEAE 379

Leuc. gasicomitatum (AB354939)

Leuc. gelidum (AB354941)
Leuc. inhae (AB354947)
Leuc. citreum (AB354943)
Leuc. kimchii (CP001758)
Leuc. carnosum (AB354949)
Leuc. lactis (AB354945)
Leuc. holzapfelii (AM940229)
Leuc. palmae (AM940226)
Leuc. pseudomesenteroides (AB354942)
Leuc. fallax (AB354948)
Leuc. mesenteroides (AB354944)
Fru. durionis (AB354950)
Fru. pseudoficulneus (AB354951)
Fru. fructosus (AB354946)
Fru . ficulneus (AB354940)
Fru. tropaeoli (AB542055)
O. oeni (CP000411)
W. viridescens (AB354952)
W. minor (J621693)
W. confusa (AB354954)
W. paramesenteroides (AB354953)
W. kandleri (AJ621692)
Lc. lactis (AE006272)
Strep. pyogenes (U21934)
Ped. acidilactici (GG730086)
Ped. pentosaceus (AJ621699)
Lb. kunkeei (AJ621652)
Lb. sakei (AJ621674)
Lb. casei (AJ621627)
Lb. plantarum (AL935258)
Lb. delbrueckii (CR954253)
Bacillus subtilis (AP011541)
0.1

Figure V.2 Phylogenetic relationships among the family Leuconostocaceae and related taxa based on recA gene sequences. The tree was
constructed by the neighbour-joining method. Bacillus subtilis was used as an outgroup. Bootstrap values are given at branching points;
values above 70% are indicated

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