Candidatus Liberibacter
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''Liberibacter'' is a genus of
Gram-negative bacteria Gram-negative bacteria are bacteria that, unlike gram-positive bacteria, do not retain the Crystal violet, crystal violet stain used in the Gram staining method of bacterial differentiation. Their defining characteristic is that their cell envelo ...
in the
Rhizobiaceae The Rhizobiaceae is a family of Pseudomonadota comprising multiple subgroups that enhance and hinder plant development. Some bacteria found in the family are used for plant nutrition and collectively make up the rhizobia. Other bacteria such as ' ...
family. Detection of the liberibacteria is based on PCR amplification of their
16S rRNA gene 16S ribosomal RNA (or 16 S rRNA) is the RNA component of the 30S subunit of a prokaryotic ribosome (SSU rRNA). It binds to the Shine-Dalgarno sequence and provides most of the SSU structure. The genes coding for it are referred to as 16S rRN ...
with specific primers. Members of the genus are plant pathogens mostly transmitted by
psyllids Psyllidae, the jumping plant lice or psyllids, are a Family (biology), family of small plant-feeding insects that tend to be very host-specific, i.e. each plant-louse species only feeds on one plant species (monophagous) or feeds on a few closel ...
. The genus was originally spelled ''Liberobacter''. Most importantly, ''Liberibacter'' is a causative agent of Huanglongbing disease (HLB) also known as citrus greening disease. ''Liberibacter'' is transmitted by two insects from
Psyllidae Psyllidae, the jumping plant lice or psyllids, are a Family (biology), family of small plant-feeding insects that tend to be very host-specific, i.e. each plant-louse species only feeds on one plant species (monophagous) or feeds on a few closel ...
family – ''
Diaphorina citri ''Diaphorina citri'', the Asian citrus psyllid, is a sap-sucking, hemipteran bug now in the taxonomic family Liviidae. It is one of two confirmed vectors of citrus greening disease. It has a wide distribution in southern Asia and has spread to ...
'' in Asia, Brazil and Florida, and ''
Trioza erytreae ''Trioza erytreae'', the African citrus psyllid, is a sap-sucking insect, a hemipteran bug in the family Triozidae. It is an important pest of citrus, being one of only two known vectors of the serious citrus disease, '' huanglongbing'' or citru ...
'' in Africa. The Asian HLB strain, "''Candidatus'' Liberibacter asiaticus" is more heat tolerant, while the African strain, ''Candidatus'' Liberibacter africanus is asymptomatic at temperatures above 30 °C. Species of ''Liberibacter'', infecting solanaceous plants has been identified and it was carried by another psyllid, a potato pest ''Bactericera cockerelli''.


Genomes

The genetic diversity within the genus is best expressed as the diversity across
genomes A genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding genes, other functional regions of the genome such as ...
. More than 60 genomes have been sequenced, ranging in size from 233 kb to about 1.5 MB, hence the genomes are small compared to most other bacteria. The smallest genome (Candidatus ''Liberibacter asiaticus'' strain SGCA1) encodes only 655 proteins, while the largest genome (Candidatus ''Liberibacter asiaticus'' Tabriz. 3) encodes 2174 proteins. The small genome size is typical for pathogenic bacteria which often undergo
genome reduction Genome size is the total amount of DNA contained within one copy of a single complete genome. It is typically measured in terms of mass in picograms (trillionths or 10−12 of a gram, abbreviated pg) or less frequently in daltons, or as the total ...
. This is due to adaptation to their host which often provides many nutrients, so that the
parasite Parasitism is a Symbiosis, close relationship between species, where one organism, the parasite, lives (at least some of the time) on or inside another organism, the Host (biology), host, causing it some harm, and is Adaptation, adapted str ...
does not need genes to produce those nutrients itself.


Pathogenicity

''Liberibacter'' bacteria are carried in the
hemolymph Hemolymph, or haemolymph, is a fluid, similar to the blood in invertebrates, that circulates in the inside of the arthropod's body, remaining in direct contact with the animal's tissues. It is composed of a fluid plasma in which hemolymph c ...
and
salivary gland The salivary glands in many vertebrates including mammals are exocrine glands that produce saliva through a system of ducts. Humans have three paired major salivary glands ( parotid, submandibular, and sublingual), as well as hundreds of min ...
s of psyllids. Since psyllids feed on
sap Sap is a fluid transported in the xylem cells (vessel elements or tracheids) or phloem sieve tube elements of a plant. These cells transport water and nutrients throughout the plant. Sap is distinct from latex, resin, or cell sap; it is a s ...
, this provides bacteria the entry to
phloem Phloem (, ) is the living tissue in vascular plants that transports the soluble organic compounds made during photosynthesis and known as ''photosynthates'', in particular the sugar sucrose, to the rest of the plant. This transport process is ...
of the plant. They induce significant metabolic and regulatory changes that damage the plants transport system and affects plants defense systems. These impairments have downstream negative effects on citrus microbiome of the infected plants. Since pathological ''Liberibacter'' cannot be cultivated outside of its vector or host, genetics, bacteria-vector and bacteria-plant interaction have not yet been thoroughly explored. Factors important for adaption and colonization or possible coevolution are not yet understood. Only in 2014 the accidental discovery of ''Liberibacter crescens'' in
Babaco The babaco (''Vasconcellea'' × ''heilbornii''; syn. ''Carica pentagona''), is a hybrid cultivar in the genus '' Vasconcellea'' from Ecuador. It is a hybrid between '' Vasconcellea cundinamarcensis'' (syn. ''Carica pubescens'', "mountain papaya ...
papaya The papaya (, ), papaw, () or pawpaw () is the plant species ''Carica papaya'', one of the 21 accepted species in the genus '' Carica'' of the family Caricaceae, and also the name of its fruit. It was first domesticated in Mesoamerica, within ...
(during a Papaya Bunchy Top Disease study), which can be cultured axenically, allowed researchers to establish the
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
and provided a valuable model organism to study related HLB strains. ''Liberibacter'' activates salicylic acid pathway in host, likely due to recognition of extracellular molecules such as lipopolyscacharides or
flagella A flagellum (; : flagella) (Latin for 'whip' or 'scourge') is a hair-like appendage that protrudes from certain plant and animal sperm cells, from fungal spores ( zoospores), and from a wide range of microorganisms to provide motility. Many pr ...
. Pathogen in turn likely mitigates the effects, because it encodes SA hydroxylase, that degrades
salicylic acid Salicylic acid is an organic compound with the formula HOC6H4COOH. A colorless (or white), bitter-tasting solid, it is a precursor to and a active metabolite, metabolite of acetylsalicylic acid (aspirin). It is a plant hormone, and has been lis ...
. Liberibacteria were shown to affect the spread of vector, by influencing the flight frequencies and sexual attraction of ''D. citri''. On the other hand, infection with ''Liberibacter'' causes higher mortality of ''D. citri'' adults, but not nymphs. ''Liberibacter'' is a part of the psyllid microbiota and co-existence with other bacteria likely has impact on the overall fitness of the insect, as well as outcome of the disease.


Treatment

Primary strategy for HLB disease management is a vector control. Antimicrobial treatment can suppress ''Liberibacter'' species, however usage of broad spectrum antibiotics is inadvisable due to adverse environmental effects. Alternative treatments, such as heat therapy, ''i.e.'', incubation of plant at temperatures above 40 °C for several days, show varying effects. Another suggested alternatives include the use of compounds that alleviate disease symptoms and boost plants defense systems or reinforcing natural citrus microbiota in order to compete with ''Liberibacter'' species. Early detection of HLB positive trees and removal from the groves, and extensive control of psyllids are the crucial HLB management strategies.


Species

Named species include: * " ''Ca.'' Liberibacter africanus" corrig. Jagoueix ''et al''. 1994 originated in Africa and is a causal agent of
citrus greening disease Citrus greening disease ( abbr. HLB) is a disease of citrus caused by a vector-transmitted pathogen. The causative agents are motile bacteria, '' Liberibacter'' spp. The disease is transmitted by the Asian citrus psyllid, '' Diaphorina citri'', ...
, also known as huanglongbing, and vectored by the African citrus psyllid ''
Trioza erytreae ''Trioza erytreae'', the African citrus psyllid, is a sap-sucking insect, a hemipteran bug in the family Triozidae. It is an important pest of citrus, being one of only two known vectors of the serious citrus disease, '' huanglongbing'' or citru ...
''. * " ''Ca.'' Liberibacter americanus" Teixeira ''et al''. 2005 is a novel species from Brazil described in 2005 and associated with huanglongbing and vectored by the Asian citrus psyllid ''
Diaphorina citri ''Diaphorina citri'', the Asian citrus psyllid, is a sap-sucking, hemipteran bug now in the taxonomic family Liviidae. It is one of two confirmed vectors of citrus greening disease. It has a wide distribution in southern Asia and has spread to ...
''. * " ''Ca.'' Liberibacter asiaticus" corrig. Jagoueix ''et al''. 1994 originated in Asia and is a causal agent of huanglongbing, vectored by the Asian citrus psyllid ''D. citri''. * " ''Ca.'' Liberibacter brunswickensis" Morris ''et al''. 2017 associated with the psyllid '' Acizzia solanicola'' on eggplant in Australia. * '' Liberibacter crescens'' Fagen ''et al''. 2014 was isolated from papaya growing in Puerto Rico. * " ''Ca.'' Liberibacter europaeus" Raddadi ''et al''. 2011 is a novel species described in 2010, found in
pear Pears are fruits produced and consumed around the world, growing on a tree and harvested in late summer into mid-autumn. The pear tree and shrub are a species of genus ''Pyrus'' , in the Family (biology), family Rosaceae, bearing the Pome, po ...
trees, where it seems to cause no symptoms and is vectored by the
psyllid Psyllidae, the jumping plant lice or psyllids, are a family of small plant-feeding insects that tend to be very host-specific, i.e. each plant-louse species only feeds on one plant species (monophagous) or feeds on a few closely related plants ( ...
, ''
Cacopsylla pyri ''Cacopsylla pyri'', commonly known as the pear psylla, pear psyllid, or by its synonym ''Psylla pyri'', is an insect in the subfamily Psyllinae. Originating in Europe and Asia, it has spread to North America. It is a pest of pear trees, sucking ...
''. * " ''Ca.'' Liberibacter solanacearum" Liefting ''et al''. 2009 is a causal agent of
zebra chip Zebra chip, also known as ''papa manchada'' and ''papa rayada'', is a disease in potatoes putatively caused by an alphaproteobacterium "''Candidatus'' Liberibacter solanacearum", which is vectored by the potato psyllid. When fried, potato tubers ...
disease in potatoes. It can also infect other economically important crops including tomatoes, carrot, parsely, parsnip, celery and chervil. There are at least ten
haplotypes A haplotype (haploid genotype) is a group of alleles in an organism that are inherited together from a single parent. Many organisms contain genetic material (DNA) which is inherited from two parents. Normally these organisms have their DNA orga ...
described within this species, designated LsoA, LsoB, LsoC, LsoD, LsoE, Lso F, LsoG, LsoH, LsoH(Con) and LsoU. Haplotypes A, B and F are associated with
solanaceous Solanaceae (), commonly known as the nightshades, is a family of flowering plants in the order Solanales. It contains approximately 2,700 species, several of which are used as agricultural crops, medicinal plants, and ornamental plants. Many mem ...
plants (potatoes and tomato) and vectored by the potato tomato psyllid ''
Bactericera cockerelli ''Bactericera cockerelli'', also known as the potato psyllid, is a species of Psylloidea, psyllid native to southern North America. Its range extends from Central America north to the American Pacific Northwest and parts of Manitoba, in Canada ...
''. Haplotypes C, D, E and H affect apiaceous crops (carrots, celery etc). Haplotypes D and E are vectored by ''Bactericera trigonica''. Haplotype C is vectored by ''Trioza apicalis.'' The vector for haplotype H is currently unknown. Haplotype U has been found in ''
Urtica dioica ''Urtica dioica'', often known as common nettle, burn nettle, stinging nettle (although not all plants of this species sting) or nettle leaf, or just a nettle or stinger, is a herbaceous perennial flowering plant in the family Urticaceae. Or ...
'' (stinging nettle) and is vectored by''Trioza urticae.'' ** LsoA is also described as a species as " ''Ca.'' Liberibacter psyllidaureus" corrig. Hansen ''et al''. 2008 or the misspelling ''Ca.'' L. psyllaurous. It is now considered synonymous as the 16S rRNA genes are identical. In addition to being a plant pathogen, LsoA also serves as an endosymbiont for the insect vector, by modifying tomato defenses in favor of itself and its vector. * "''Ca.'' Liberibacter ctenarytainae" is vectored by '' Ctenarytaina fuchsiae''. Its genome was announced under a different spelling, "''Ca.'' Liberibacter ctenarytaina". * "''Ca.'' Liberibacter capsica" is vectoed by '' Russelliana capsici''.


References


Further reading

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External links


g__Liberibacter
in
GTDB The Genome Taxonomy Database (GTDB) is an online database that maintains information on a proposed nomenclature of prokaryotes, following a phylogenomic approach based on a set of conserved single-copy proteins. In addition to resolving paraphyl ...
{{Taxonbar, from=Q4262259 Bacterial plant pathogens and diseases Potato diseases Rhizobiaceae