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Plant responsiveness to mycorrhizas differs from dependence upon mycorrhizas.

Soil phosphorus response curves of plants with and without mycorrhizas reflect two different, but complementary, phenomena. The first, plant responsiveness to mycorrhizas, is represented by the difference in growth between plants with and without mycorrhizas at any designated level of phosphorus availability. This is also a measure of mycorrhizal fungus effectiveness. The second, the lowest level of phosphorus availability at which plants can grow without mycorrhizas, is here termed dependence upon mycorrhizas. The latter definition differs from conventional usage which fails to distinguish dependence from responsiveness. Sigmoid curves generated by the three-parameter, logistic equation generally can model the responses of plants to mycorrhizas and phosphorus addition and can be used to assess responsiveness, effectiveness, and dependence. Such curves reveal that plant responsiveness or fungus effectiveness determined at a single level of phosphorus availability may be misleading when used to compare different host species' intrinsic capacities to respond to different mycorrhizal fungus species. Instead, the same relative position should be evaluated among phosphorus response curves for different species combinations. Dependence of a plant species known to benefit from mycorrhizas can be assessed with reference to only the phosphorus response curve of plants without mycorrhizas. Dependence is a constitutive property of plant species that can be used to classify them as facultatively or obligately mycotrophic. Dependence is a plant attribute upon which natural selection can act, but responsiveness and effectiveness cannot be selected directly because they are emergent properties of the interaction between plant and fungus species.

Models, Biological↗

Mycorrhiza and root hairs in barley enhance acquisition of phosphorus and uranium from phosphate rock but mycorrhiza decreases root to shoot uranium transfer.

Some phosphate rocks (PR) contain high concentrations of uranium (U), which are potentially toxic via accumulation in soils and food chains, and plant uptake of U is likely to be influenced by characteristics of roots and associated microorganisms. The relative importance of root hairs and mycorrhiza in U uptake from PR was studied using a root hairless barley (Hordeum vulgare) mutant (Brb) and its wild type (WT). Both plant genotypes were grown in pots with Glomus intraradices BEG 87, or in the absence of mycorrhiza, and three P treatments were included: nil P, 2% (w/w) PR and 50 mg KH(2)PO(4)-P kg(-1) soil. Mycorrhiza markedly increased d. wts and P contents of Brb amended with nil P or PR, but generally depressed d. wts of WT plants, irrespective of P amendments. Mycorrhiza had contrasting effects on U contents in roots and shoots, in particular in Brb where mycorrhiza increased root U concentrations but decreased U translocation from roots to shoots. The experiment supports our understanding of arbuscular mycorrhiza as being multifunctional by not only improving the utilization of PR by the host plant but also by contributing to the phytostabilization of uranium.

Hordeum↗

Expression of early nodulin genes in alfalfa mycorrhizae indicates that signal transduction pathways used in forming arbuscular mycorrhizae and Rhizobium-induced nodules may be conserved.

Transcripts for two genes expressed early in alfalfa nodule development (MsENOD40 and MsENOD2) are found in mycorrhizal roots, but not in noncolonized roots or in roots infected with the fungal pathogen Rhizoctonia solani. These same two early nodulin genes are expressed in uninoculated roots upon application of the cytokinin 6-benzylaminopurine. Correlated with the expression of the two early nodulin genes, we found that mycorrhizal roots contain higher levels of trans-zeatin riboside than nonmycorrhizal roots. These data suggest that there may be conservation of signal transduction pathways between the two symbioses-nitrogen-fixing nodules and phosphate-acquiring mycorrhizae.

Journal Article↗

The influence of mycorrhiza on uranium and phosphorus uptake by barley plants from a field-contaminated soil.

BACKGROUND: Recent studies indicated that arbuscular mycorrhizal fungi (AMF) play important roles in plant accumulation of uranium (U) from contaminated environments, but the impacts of fertilization practices on functioning of the symbiotic associations, which are crucial factors influencing plant nutrition and growth responses to mycorrhiza, have rarely been considered. MATERIALS AND METHODS: In a greenhouse experiment, a bald root barley mutant (brb) together with the wild type (wt) were used to test the role of root hairs and AMF in uranium (U) uptake by host plants from a U contaminated soil. Nil, 20 and 60 mg KH2PO4-P kg(-1) soil were included to investigate the influences of phosphorus (P) fertilization on plant growth and accumulation of U. RESULTS: Dry matter yield of barley plants increased with increasing P additions and wt produced significantly higher dry weight than brb. Mycorrhiza markedly improved dry matter yield of both genotypes grown at nil P, whereas only brb responded positively to mycorrhiza at 20 mg P kg(-1). At the highest P level, mycorrhiza resulted in growth depressions in both genotypes, except for the roots of wt. In general, plant P concentrations increased markedly with increasing P additions and in response to mycorrhiza. Mycorrhiza and P additions had no significant effects on shoot U concentrations. However, root U concentrations in both genotypes were significantly increased by mycorrhiza. On the other hand, shoot U contents increased with increasing P levels, while 20 mg P kg(-1) stimulated, but 60 mg P kg(-1) marginally affected the U accumulation in roots. Root length specific U uptake was moderately enhanced both by root hairs and strongly enhanced by mycorrhiza. Moreover, non-inoculated plants generally had higher shoot-root ratios of U content than the corresponding inoculated controls. CONCLUSION: Our study shows that AMF and root hairs improves not only P acquisition but also the root uptake of U, and mycorrhiza generally decreases U translocation from plant root to shoot. Hence, mycorrhiza is of potential use in the phytostabilization of U contaminated environments. PERSPECTIVES: The complex impacts of P on U accumulation by barley plants suggested that U behavior in mycorrhizosphere and translocation along the soil-fungi-plant continuum as affected by fertilization practices deserve extensive studies for optimizing the function of mycorrhizal associations for phytoremediation purposes.

Biodegradation, Environmental↗

Morphological characterization of the mycorrhiza formed by Helianthemum almeriense Pau with Terfezia claveryi Chatin and Picoa lefebvrei (Pat.) Maire.

This work presents the first anatomical description of the mycorrhizal systems of Helianthemum almeriense, and of the structure and ultrastructure of the mycorrhizae formed by this plant species with the ascomycetes Terfezia claveryi and Picoa lefebvrei. Four different mycorrhizal systems are described, the club-shaped mycorrhiza being the most abundant. The type of mycorrhiza formed depended on the mycorrhiza culture conditions, but not on the fungal species. For both fungal species, H. almeriense formed an endomycorrhiza in natural field conditions, an ecto- and ectendomycorrhiza without a sheath in pot cultures, and an ectomycorrhiza with a characteristic sheath and Hartig net in in vitro cultures. This is the first report of a typical sheath in Helianthemum-desert truffle mycorrhizal associations. The results support the idea that culture conditions can induce changes in mycorrhiza morphology and that there is no clear barrier between the two main types of mycorrhiza organization in Helianthemum species. The ultrastructural study confirmed the regular presence of T. claveryi intracellular hyphae in direct contact with the host wall, a localization which seems to be a characteristic of the T. claveryi mycorrhiza organization. The P. lefebvrei mycorrhiza organization was characterized by intracellular hyphae with large amounts of electron-dense globules, probably with a lipidic content, and a warty ornamentation on the wall of the root external hyphae.

Ascomycota↗

Epiphytic and terrestrial mycorrhizas in a lower montane Costa Rican cloud forest.

The epiphyte community is the most diverse plant community in neotropical cloud forests and its collective biomass can exceed that of the terrestrial shrubs and herbs. However, little is known about the role of mycorrhizas in this community. We assessed the mycorrhizal status of epiphytic (Araceae, Clusiaceae, Ericaceae, and Piperaceae) and terrestrial (Clusiaceae, Ericaceae) plants in a lower montane cloud forest in Costa Rica. Arbuscular mycorrhizas were observed in taxa from Araceae and Clusiaceae; ericoid mycorrhizas were observed in ericaceous plants. This is the first report of intracellular hyphal coils characteristic of ericoid mycorrhizas in roots of Cavendishia melastomoides, Disterigma humboldtii, and Gaultheria erecta. Ericaceous roots were also covered by an intermittent hyphal mantle that penetrated between epidermal cells. Mantles, observed uniquely on ericaceous roots, were more abundant on terrestrial than on epiphytic roots. Mantle abundance was negatively correlated with gravimetric soil water content for epiphytic samples. Dark septate endophytic (DSE) fungi colonized roots of all four families. For the common epiphyte D. humboldtii, DSE structures were most abundant on samples collected from exposed microsites in the canopy. The presence of mycorrhizas in all epiphytes except Peperomia sp. suggests that inoculum levels and environmental conditions in the canopy of tropical cloud forests are generally conducive to the formation of mycorrhizas. These may impact nutrient and water dynamics in arboreal ecosystems.

Araceae↗

Co-occurrence of Arum- and Paris-type morphologies of arbuscular mycorrhizae in cucumber and tomato.

Colonization by arbuscular mycorrhizal (AM) fungi was investigated in cucumber (Cucumis sativus), tomato (Lycopersicon esculentum) and Clethra barbinervis (Ericales) grown in field-collected soil known from previous studies to generate Paris-type arbuscular mycorrhizae in C. barbinervis. Spores of Paraglomus, Acaulospora, Glomus, and Gigaspora were found in the soil. Formation of hyphal coils and arbusculate coils of Paris-type mycorrhizae and of arbuscules of Arum-type mycorrhizae in roots raised in this soil in the growth chamber were compared with the detection of DNA of AM fungi from the same root systems using Glomales-specific primers. Only Paris-type mycorrhizae with extensive arbusculate coils developed in C. barbinervis, but cucumber and tomato developed both Paris- and Arum-types in the same root systems. Glomaceae and Archaeosporaceae and/or Paraglomaceae were detected strongly in the DNA from both cucumber and tomato roots, in which Arum-type mycorrhizae were observed. In contrast, DNA of Glomaceae was detected more sparingly in C. barbinervis, in which Paris-type mycorrhizae dominated. Acaulosporaceae and Gigasporaceae were strongly detected in the DNA from both C. barbinervis and tomato, whereas they were more weakly detected in cucumber. These results indicate that the morphology of colonization is strongly influenced by the selection of fungi to colonize the host plant from among those in the soil environment.

Clethraceae↗

The role of arbuscular mycorrhiza on change of heavy metal speciation in rhizosphere of maize in wastewater irrigated agriculture soil.

To understand the roles of mycorrhiza in metal speciation in the rhizosphere and the impact on increasing host plant tolerance against excessive heavy metals in soil, maize (Zea mays L.) inoculated with arbuscular mycorrhizal fungus (Glomus mosseae) was cultivated in heavy metal contaminated soil. Speciations of copper, zinc and lead in the soil were analyzed with the technique of sequential extraction. The results showed that, in comparison to the bolked soil, the exchangeable copper increased from 26% to 43% in non-infected and AM-infected rhizoshpere respectively; while other speciation (organic, carbonate and Fe-Mn oxide copper) remained constant and the organic bound zinc and lead also increased but the exchangeable zinc and lead were undetectable. The organic bound copper, zinc and lead were higher by 15%, 40% and 20%, respectively, in the rhizosphere of arbuscular mycorrhiza infected maize in comparison to the non-infected maize. The results might indicate that mycorrhiza could protect its host plants from the phytotoxicity of excessive copper, zinc and lead by changing the speciation from bio-available to the non-bio-available form. The fact that copper and zinc accumulation in the roots and shoots of mycorrhia infected plants were significantly lower than those in the non-infected plants might also suggest that mycorrhiza efficiently restricted excessive copper and zinc absorptions into the host plants. Compared to the non-infected seedlings, the lead content of infected seedlings was 60% higher in shoots. This might illustrate that mycorrhiza have a different mechanism for protecting its host from excessive lead phytotoxicity by chelating lead in the shoots.

Analysis of Variance↗

Uptake and speciation of selenium in garlic cultivated in soil amended with symbiotic fungi (mycorrhiza) and selenate.

The scope of the work was to investigate the influence of selenate fertilisation and the addition of symbiotic fungi (mycorrhiza) to soil on selenium and selenium species concentrations in garlic. The selenium species were extracted from garlic cultivated in experimental plots by proteolytic enzymes, which ensured liberation of selenium species contained in peptides or proteins. Separate extractions using an aqueous solution of enzyme-deactivating hydroxylamine hydrochloride counteracted the possible degradation of labile selenium species by enzymes (such as alliinase) that occur naturally in garlic. The selenium content in garlic, which was analysed by ICP-MS, showed that addition of mycorrhiza to the natural soil increased the selenium uptake by garlic tenfold to 15 microg g(-1) (dry mass). Fertilisation with selenate and addition of mycorrhiza strongly increased the selenium content in garlic to around one part per thousand. The parallel analysis of the sample extracts by cation exchange and reversed-phase HPLC with ICP-MS detection showed that gamma-glutamyl-Se-methyl-selenocysteine amounted to 2/3, whereas methylselenocysteine, selenomethionine and selenate each amounted to a few percent of the total chromatographed selenium in all garlic samples. Se-allyl-selenocysteine and Se-propyl-selenocysteine, which are selenium analogues of biologically active sulfur-containing amino acids known to occur in garlic, were searched for but not detected in any of the extracts. The amendment of soil by mycorrhiza and/or by selenate increased the content of selenium but not the distribution of detected selenium species in garlic. Finally, the use of two-dimensional HPLC (size exclusion followed by reversed-phase) allowed the structural characterisation of gamma-glutamyl-Se-methyl-selenocysteine and gamma-glutamyl-Se-methyl-selenomethionine in isolated chromatographic fractions by quadrupole time-of-flight mass spectrometry.

Enzymes↗

The conserved arbuscular mycorrhiza-specific transcription of the secretory lectin MtLec5 is mediated by a short upstream sequence containing specific protein binding sites.

In Medicago truncatula a family of mycorrhiza-specific expressed lectins has been identified recently, but the function and regulation of these lectins during the arbuscular mycorrhiza symbiosis are still unknown. In order to characterize a first member of this protein family, MtLec5 was analyzed concerning its localization and regulation. Confocal laser scanning microscopy showed that MtLec5 is a secretory protein indicating a role as a vegetative storage protein, which is specifically expressed in mycorrhizal root systems. To study the molecular mechanisms leading to the mycorrhiza-specific transcription, deletion studies of pMtLec5 were done using reporter gene fusions. Potential cis-acting elements could be narrowed down to a 150 bp fragment that was located approximately at -300/-150 according to the transcription start, suggesting the binding of positive regulators to this area. Similar expression pattern of the reporter gene was found after transforming roots of the non-legume Nicotiana tabacum with the heterologous promoter-reporter fusions. This indicated that the observed mycorrhiza-specific transcriptional induction is not legume-specific. Electrophoretic mobility shift assays showed that several factors which were exclusively present in mycorrhizal roots bind within the 150 bp promoter area. This strengthens the hypothesis of positive regulators mediating the AM-specific gene expression.

Base Sequence↗

Arbuscular mycorrhiza and Collembola interact in affecting community composition of saprotrophic microfungi.

The functioning of the plant-mycorrhiza system depends on interactions with other organisms, including saprotrophic (ST) soil fungi. The interactions between mycorrhizal and ST fungi are likely affected by fungivorous soil animals, such as Collembola. In a two-factorial laboratory experiment lasting for 30 weeks we assessed the effects of an arbuscular mycorrhizal fungus (Glomus mosseae) and Collembola (Protaphorura fimata, Heteromurus nitidus and Folsomia candida) on the community composition of ST microfungi in soil planted with the invasive grass Cynodon dactylon. The presence of mycorrhiza substantially reduced total plant biomass and reduced N and P availability to the soil microflora, though these effects were less pronounced in the presence of Collembola. The density of Collembola was high (corresponding to about 2x10(5) individuals m-2) and was not affected by the presence of G. mosseae. In spite of the large amount of mycorrhizal mycelium in soil, it contributed little to Collembola nutrition. The presence of mycorrhiza strongly affected the community structure of ST soil fungi. In particular, mycorrhiza reduced the relative abundance of Trichoderma harzianum and Exophiala sp., but increased the abundance of Ramichloridium schulzeri and several sterile forms. However, the difference between fungal communities in mycorrhizal and non-mycorrhizal treatments was much more pronounced in the presence of Collembola. Presumably, the intense grazing by Collembola destabilized the ST fungal community, thereby making it more susceptible to the influence of G. mosseae. These results document for the first time that fungal feeding soil invertebrates can significantly affect the interactions between mycorrhizal fungi and ST soil microorganisms.

Animals↗

Methods to estimate the proportion of plant and fungal RNA in an arbuscular mycorrhiza.

Arbuscular mycorrhizas are endosymbiotic associations formed between obligately biotrophic arbuscular mycorrhizal (AM) fungi and plant roots. The fungus and plant coexist in intimate contact as the fungus grows within the cortex of the root. RNA isolated from arbuscular mycorrhizas contains transcripts from both eukaryotic genomes. It is essential to be able to estimate the relative levels of fungal and plant RNA so that changes in plant and fungal gene expression can be evaluated during development of the AM symbiosis. Here we describe the design and use of specific plant and fungal internal transcribed spacer sequences and 18S rRNA probes to distinguish and quantify the relative levels of RNA of plant and fungal origin in samples from arbuscular mycorrhizas. We present two different methods. The first employs the most traditional method of transcript level analysis, namely northern blot analysis. The second one uses ribonuclease protection assays, which permit the analysis of transcript levels in a very small amount of tissue and are proving to be suitable for the analysis of gene expression in AM fungi. Analysis of tissues from a developing mycorrhiza showed that the relative levels of fungal RNA increased gradually as colonization of the root system progressed, reaching 5-12% in the most highly colonized samples.

Agaricales↗

Mycorrhiza-plant colonization patterns on a subalpine glacier forefront as a model system of primary succession.

Lyman glacier in the North Cascades Mountains of Washington has a subalpine forefront characterized by a well-developed terminal moraine, inconspicuous successional moraines, fluting, and outwash. These deposits were depleted of symbiotic fungi when first exposed but colonized by them over time after exposure. Four major groups of plant species in this system are (1) mycorrhiza-independent or facultative mycotrophic, (2) dependent on arbuscular mycorrhizae (AM) (3) dependent on ericoid mycorrhiza (ERM) or ectomycorrhizae (EM), and (4) colonized by dark-septate (DS) endophytes. We hypothesized that availability of mycorrhizal propagules was related to the success of mycorrhiza-dependent plants in colonizing new substrates in naturally evolved ecosystems. To test this hypothesis roots samples of 66 plant species were examined for mycorrhizal colonization. The plants were sampled from communities at increasing distances from the glacier terminus to compare the newest communities with successively older ones. Long established, secondary successional dry meadow communities adjacent to the glacier forefront, and nearby high alpine communities were sampled for comparison. DS were common on most plant species on the forefront. Nonmycorrhizal plants predominated in the earlier successional sites, whereas the proportion of mycorrhizal plants generally increased with age of community. AM were present, mostly at low levels, and nearly absent in two sites of the forefront. ERM were present in all species of Ericaceae sampled, and EM in all species of Pinaceae and Salicaceae. Roots of plants in the long established meadow and heath communities adjacent to the forefront and the high alpine community all had one or another of the colonization types, with DS and AM predominating.

Ecosystem↗

Arbuscular mycorrhiza reduces susceptibility of tomato to Alternaria solani.

Mycorrhiza frequently leads to the control of root pathogens, but appears to have the opposite effect on leaf pathogens. In this study, we studied mycorrhizal effects on the development of early blight in tomato (Solanum lycopersicum) caused by the necrotrophic fungus Alternaria solani. Alternaria-induced necrosis and chlorosis of all leaves were studied in mycorrhizal and non-mycorrhizal plants over time course and at different soil P levels. Mycorrhizal tomato plants had significantly less A. solani symptoms than non-mycorrhizal plants, but neither plant growth nor phosphate uptake was enhanced by mycorrhizas. An increased P supply had no effect on disease severity in non-mycorrhizal plants, but led to a higher disease severity in mycorrhizal plants. This was parallel to a P-supply-induced reduction in mycorrhiza formation. The protective effect of mycorrhizas towards development of A. solani has some parallels to induced systemic resistance, mediated by rhizobacteria: both biocontrol agents are root-associated organisms and both are effective against necrotrophic pathogens. The possible mechanisms involved are discussed.

Alternaria↗

Identification of mycorrhiza-regulated genes with arbuscule development-related expression profile.

Suppressive subtractive hybridisation was applied to the analysis of late stage arbuscular mycorrhizal development in pea. 96 cDNA clones were amplified and 81, which carried fragments more than 200 nt in size, were sequence analysed. Among 67 unique fragments, 10 showed no homology and 10 were similar to sequences with unknown function. RNA accumulation of the corresponding 67 genes was analysed by hybridisation of macro-arrays. The cDNAs used as probes were derived from roots of wild type and late mutant pea genotypes, inoculated or not with the AM fungus Glomus mosseae. After calibration, a more than 2.5-fold mycorrhiza-induced RNA accumulation was detected in two independent experiments in the wild type for 25 genes, 22 of which seemed to be induced specifically during late stage AM development. Differential expression for 7 genes was confirmed by RT-PCR using RNA from mycorrhiza and from controls of a different pea cultivar. In order to confirm arbuscule-related expression, the Medicago truncatula EST data base was screened for homologous sequences with putative mycorrhiza-related expression and among a number of sequences with significant similarities, a family of trypsin inhibitor genes could be identified. Mycorrhiza-induced RNA accumulation was verified for five members by real-time PCR and arbuscule-related activation of the promoter could be shown in transgenic roots for one of the genes, MtTi 1.

Expressed Sequence Tags↗

Arbuscular mycorrhizae enhance metal lead uptake and growth of host plants under a sand culture experiment.

A sand culture experiment was conducted to investigate whether mycorrhizal colonization and mycorrhizal fungal vesicular numbers were influenced by metal lead, and whether mycorrhizae enhance host plants tolerance to metal lead. Metal lead was applied as Pb(NO3)2 in solution at three levels (0, 300 and 600 mg kg(-1) sand). Five mycorrhizal host plant species, Kummerowia striata (Thunb.) Schindl, Ixeris denticulate L., Lolium perenne L., Trifolium repens L. and Echinochloa crusgalli var. mitis were used to examine Pb-mycorrhizal interactions. The arbuscular mycorrhizal inoculum consisted of mixed spores of mycorrhizal fungal species directly isolated from orchard soil. Compared to the untreated control, both Pb concentrations reduced mycorrhizal colonization by 3.8-70.4%. Numbers of AM fungal vesicles increased by 13.2-51.5% in 300 mg Pb kg(-1) sand but decreased by 9.4-50.9% in 600 mg Pb kg(-1) sand. Mycorrhizae significantly enhanced Pb accumulation both in shoot by 10.2-85.5% and in root by 9.3-118.4%. Mycorrhizae also enhanced shoot biomass and shoot P concentration under both Pb concentrations. Root/shoot ratios of Pb concentration were higher in highly mycorrhizal plant species (K.striata, I. denticulate, and E. crusgalli var. mitis) than that in poorly mycorrhizal ones (L. perenne and T. repens,). Mycorrhizal inoculation increased the root/shoot ratio of Pb concentration of highly mycorrhizal plant species by 7.6-57.2% but did not affect the poorly mycorrhizal ones. In the treatments with 300 Pb mg kg(-1) sand, plant species with higher vesicular numbers tended to show higher root/shoot ratios of the Pb concentration. We suggest that under an elevated Pb condition, mycorrhizae could promote plant growth by increasing P uptake and mitigate Pb toxicity by sequestrating more Pb in roots.

Lead↗

The characterization of novel mycorrhiza-specific phosphate transporters from Lycopersicon esculentum and Solanum tuberosum uncovers functional redundancy in symbiotic phosphate transport in solanaceous species.

Solanaceous species are among the >200 000 plant species worldwide forming a mycorrhiza, that is, a root living in symbiosis with soil-borne arbuscular-mycorrhizal (AM) fungi. An important parameter of this symbiosis, which is vital for ecosystem productivity, agriculture, and horticulture, is the transfer of phosphate (Pi) from the AM fungus to the plant, facilitated by plasma membrane-spanning Pi transporter proteins. The first mycorrhiza-specific plant Pi transporter to be identified, was StPT3 from potato [Nature414 (2004) 462]. Here, we describe novel Pi transporters from the solanaceous species tomato, LePT4, and its orthologue StPT4 from potato, both being members of the Pht1 family of plant Pi transporters. Phylogenetic tree analysis demonstrates clustering of both LePT4 and StPT4 with the mycorrhiza-specific Pi transporter from Medicago truncatula [Plant Cell, 14 (2002) 2413] and rice [Proc. Natl Acad. Sci. USA, 99 (2002) 13324], respectively, but not with StPT3, indicating that two non-orthologous mycorrhiza-responsive genes encoding Pi transporters are co-expressed in the Solanaceae. The cloned promoter regions from both genes, LePT4 and StPT4, exhibit a high degree of sequence identity and were shown to direct expression exclusively in colonized cells when fused to the GUS reporter gene, in accordance with the abundance of LePT4 and StPT4 transcripts in mycorrhized roots. Furthermore, extensive sequencing of StPT4-like clones and subsequent expression analysis in potato and tomato revealed the presence of a close paralogue of StPT4 and LePT4, named StPT5 and LePT5, respectively, representing a third Pi transport system in solanaceous species which is upregulated upon AM fungal colonization of roots. Knock out of LePT4 in the tomato cv. MicroTom indicated considerable redundancy between LePT4 and other Pi transporters in tomato.

Biological Transport, Active↗