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Conflicting phylogenies of balsaminoid families and the polytomy in Ericales: combining data in a Bayesian framework.

The balsaminoid Ericales, namely Balsaminaceae, Marcgraviaceae, Tetrameristaceae, and Pellicieraceae have been confidently placed at the base of Ericales, but the relations among these families have been resolved differently in recent analyses. Sister to this basal group is a large polytomy comprising all other families of Ericales, which is associated with short internodes. Because there are more than 13 kb of sequences for a large sampling of representatives, a thorough examination of the available data with novel methods seemed in place. Because of its computational speed, Bayesian phylogenetics allows for the use of parameter-rich models that can accommodate differences in the evolutionary process between partitions in a simultaneous analysis. In addition, there are recently proposed Bayesian strategies of assessing incongruence between partitions. We have applied these methods to the current problems in Ericales phylogeny, taking into account reported pitfalls in Bayesian analysis such as model selection uncertainty. Based on our results we infer several, previously unresolved relationships in the order Ericales. In balsaminoid families, we find that the closest relatives of Balsaminaceae are Marcgraviaceae. In the Ericales polytomy, we find strong support for Pentaphylacaceae sensu APG II as the sister group of Maesaceae. In addition, Symplocaceae receive a position as sister to Theaceae and these families form a monophyletic group together with Styracaceae-Diapensiaceae. At the base of this clade are Actinidiaceae and Clethraceae. The positions of Ebenaceae and Lecythidaceae remain uncertain.

Balsaminaceae↗

The distribution and phylogeny of aluminium accumulating plants in the Ericales.

The distribution of aluminium (Al) accumulation in the Ericales is surveyed, based on semi-quantitative tests of 114 species and literature data. Al accumulation mainly characterises the families Diapensiaceae, Pentaphylacaceae, Symplocaceae, Ternstroemiaceae, and Theaceae. Al accumulation is consistently present or absent in most families examined, but the character appears to be more variable in a few taxa (e.g., Lecythidaceae, Myrsinaceae). Although the interfamilial relationships within the Ericales require further research, the ability to accumulate high levels of Al appears to show considerable taxonomic significance. While the majority of Al accumulating Ericales includes woody, tropical plants, the feature is remarkably present in several herbaceous Diapensiaceae, which have a distribution in cold to temperate areas. The association of different mycorrhizae types with plant roots is suggested to play a role in the exclusion of high Al levels from the shoot.

Aluminum↗

The micromorphology of pit membranes in tracheary elements of ericales: new records of tori or pseudo-tori?

BACKGROUND AND AIMS: Intervascular pit membranes were examined within Ericales to determine the distribution and structure of torus-like thickenings. METHODS: Forty-nine species representing 12 families of the order Ericales were investigated using light, scanning and transmission electron microscopy. They were compared with four species of Oleaceae to determine the true nature of the thickenings. KEY RESULTS: Pit membranes with torus-like thickenings were observed in seven species of Ericaceae and were found to be amorphous, plasmodesmata-associated structures with an irregular distribution. These pseudo-tori show major differences compared with true tori with respect to their distribution and ultrastructure. Genuine tori, which are strongly correlated with round pit apertures in narrow tracheary elements, were found in two species of Osmanthus (Oleaceae). CONCLUSIONS: The pseudo-tori found in some Ericaceae are considered to be similar to pit membrane thickenings previously recorded in Rosaceae. While true tori appear to be functionally significant in terms of efficiency and safety of water transport, the possible function of pseudo-tori could be associated with the role of plasmodesmata during differentiation of tracheids, fibre-tracheids or narrow vessels.

Ericaceae↗

The evolution of the atpbeta-rbcL intergenic spacer in the epacrids (Ericales) and its systematic and evolutionary implications.

Sequence data from the noncoding region separating the plastid genes atpbeta and rbcL were gathered for 27 epacrid taxa, representing all previously recognized infrafamilial groups, and four outgroup taxa (Ericaceae), to address several persistent phylogenetic questions in the group. Parsimony analyses were conducted on these data, as well as on a complementary rbcL sequence dataset assembled from the literature and the combined dataset. The atpbeta-rbcL spacer was notable for the high frequency of insertion-deletion mutations (indels); their distributions were coded as binary characters and included as a adjunct matrix in some of the analyses. The phylogenetic patterns derived from the spacer and rbcL data and the combined analyses, both including and excluding the indel data, concur in resolving seven major lineages corresponding to the tribes of Crayn et al. (1998, Aust. J. Bot. 46, 187-200), viz. Prionoteae, Archerieae, Oligarrheneae, Cosmelieae, Richeeae, Epacrideae, and Styphelieae. The relationships of the tribes and within Styphelieae, however, are not convincingly resolved. Minor conflicts in the positions of some taxa between the spacer and the rbcL trees are poorly supported. Among epacrids, the spacer region provided more cladistically informative characters than rbcL and resulted in trees with lower homoplasy. Further, the spacer data, when analyzed alone and when combined with rbcL, resolved several clades that could not be retrieved on rbcL data alone and provided increased support for many other relationships. The evolution of a putative three-base inversion associated with a hairpin secondary structure in the spacer region is discussed in the light of the inferred phylogeny.

Classification↗

Palynological variation in balsaminoid ericales. I. Marcgraviaceae.

BACKGROUND AND AIMS: Marcgraviaceae are a rather small family of seven genera and approx. 130 neotropical species. This study aims to present a detailed palynological survey of the family in order to comment on the intrafamily relationships and possible correlations with pollinators. METHODS: In total, 119 specimens representing 67 species and all genera are observed using light microscopy and scanning electron microscopy. Furthermore, eight species from five genera are studied with transmission electron microscopy. KEY RESULTS: Our results show that pollen grains of Marcgraviaceae are small (20-35 microm), have three equatorial apertures, granules on the colpus membrane, oblate spheroidal to prolate spheroidal shapes, mainly psilate to perforate ornamentations, and lalongate colpus-shaped thinnings at the inner layer of the exine, and show the presence of orbicules. Based on our fragmentary knowledge of the pollination biology of the family, there are no clear correlations between pollinators and pollen features. CONCLUSIONS: The genus Marcgravia has a high percentage of reticulate sexine patterns and a relatively thin nexine. Sarcopera can be defined by the presence of an oblate spheroidal to even suboblate shape, while Ruyschia and Souroubea typically show prolate spheroidal to subprolate pollen grains. The presence of a thick foot layer in the pollen wall is characteristic of the genera Norantea, Sarcopera and Schwartzia. Pollen features that are taxonomically useful within the family are the shape, sexine sculpturing, and ultrastructure of the pollen wall.

Genetic Variation↗

Palynological variation in Balsaminoid Ericales. II. Balsaminaceae, Tetrameristaceae, Pellicieraceae and general conclusions.

BACKGROUND AND AIMS: The objective of this study is to examine the palynological diversity of Balsaminaceae (two genera/+/-1000 species), Tetrameristaceae (two genera/two species) and Pellicieraceae (one genus/one species). The diversity found will be used to infer the systematic value of pollen features within the balsaminoid clade. METHODS: Pollen morphology and ultrastructure of 29 species, representing all families of the balsaminoid clade except Marcgraviaceae, are investigated by means of light microscopy, scanning electron microscopy and transmission electron microscopy. KEY RESULTS: Balsaminaceae pollen is small to medium sized with three to four apertures, which can be either colpate or porate, and a sexine sculpturing varying from coarsely reticulate to almost microreticulate. Tetrameristaceae pollen is small sized, 3-colporate, with a heterobrochate reticulate sculpturing and granules present in the lumina. Pellicieraceae pollen is large sized, 3-colporate with long ectocolpi and a perforate sexine sculpturing with large verrucae. Furthermore, Pelliciera is characterized by the occurrence of aggregated orbicules, while orbicules are completely absent in both Balsaminaceae and Tetrameristaceae. Balsaminaceae pollen differs from the other balsaminoid families due to the occurrence of colpate or porate grains with an oblate to peroblate shape, a very thin foot layer and a lamellated endexine. CONCLUSIONS: From a pollen morphological point of view, Balsaminaceae are completely different from the other balsaminoid families. Therefore, no pollen morphological synapomorphies could be defined for the balsaminoid clade. However, various pollen features were observed that could indicate a possible relationship between Tetrameristaceae, Pellicieraceae and Marcgraviaceae. Despite the palynological similarities in the latter three families, it remains unclear to what extent they are related to each other.

Genetic Variation↗

Commercial bumble bee Bombus impatiens (Hymenoptera: Apidae) as a pollinator in lowbush blueberry (Ericale: Ericaceae) fields.

Here, we evaluate the potential of the bumble bee Bombus impatiens Cresson, obtained from commercial colonies, as a pollen vector for lowbush blueberry, Vaccinium augustifolium Aiton. We wanted to gain insight into the relationships between pollination by introduced bumble bees, the categories of seeds produced, and the weight and the maturity of the blueberries. The effect of B. impatiens foraging in blueberry stands was measured quantitatively through fruit set and seed set analysis. We created a density gradient of B. impatiens by clustering 72 small hives at the southern extremity of blueberry fields located in Girardville (49 degrees 00' N, 72 degrees 33' W), Quebec, Canada. Observers recorded plant and insect data in 52 plots of 1 by 10 m, distributed at distances ranging from 25 to 1,500 m from the hives. From these data, we evaluated the fruit set rate, the weight and maturity of the berries, and the number of seed structures per berry, including true seeds of large, medium, and small size, pseudo-seeds, and ovules. Positive correlations were found between the density of B. impatiens and fruit set as well as the number of large seeds per berry. Large seeds influenced the weight and maturity of berries. B. impatiens acted as a "near-nest central forager" and increased fruit set and seed production up to 100 and 150 m respectively, from the hives.

Animals↗

Effects of honey bee (Hymenoptera: Apidae) and bumble bee (Hymenoptera: Apidae) presence on cranberry (Ericales: Ericaceae) pollination.

Honey bees, Apis mellifera L., are frequently used to pollinate commercial cranberries, Vaccinium macrocarpon Ait., but information is lacking on the relative contribution of honey bees and native bees, the effects of surrounding vegetation on bee visitation, and on optimal timing for honey bee introduction. We begin with a descriptive study of numbers of honey bees, bumble bees, and other bees visiting cranberry blossoms, and their subsequent effect on cranberry yield, on three cranberry properties in 1999. The property surrounded by agricultural land, as opposed to wetlands and woodlands, had fewer numbers of all bee types. In 2000, one property did not introduce honey bee colonies, providing an opportunity to document the effect of lack of honey bees on yield. With no honey bees, plants along the edge of the bed had significantly higher berry weights compared with nonedge plants, suggesting that wild pollinators were only effective along the edge. Comparing the same bed between 1999, with three honey bee colonies per acre, and 2000, with no honey bees, we found a significant reduction in average berry size. In 2000, we compared stigma loading on properties with and without honey bees. Significantly more stigmas received the minimum number of tetrads required for fruit set on the property with honey bees. Significantly more tetrads were deposited during mid-bloom compared with early bloom, indicating that mid-bloom was the best time to have honey bees present. This study emphasizes the importance and effectiveness of honey bees as pollinators of commercial size cranberry plantings.

Animals↗

Generic realignment in primuloid families of the Ericales s.l.: a phylogenetic analysis based on DNA sequences from three chloroplast genes and morphology.

The phylogenetic interrelationships in Primulaceae, Myrsinaceae, and Theophrastaceae were investigated using DNA sequence data from the chloroplast genes atpB, ndhF, and rbcL. The three genes were analyzed separately, together, and in combination with morphology, using parsimony jackknifing. The sequence data are further explored by analyses of first and second codon position only, third positions only, and transversions only. The results show that all codon positions contribute group support to the ndhF tree, whereas third codon positions provide most of the structure in the atpB and rbcL trees. Analyzed separately, transversions in atpB and rbcL have little structure, whereas in ndhF they produce a well-resolved tree. We conclude that the most informative and robust results are obtained from analyses with all codon positions included and that the tree resulting from the combined analysis of all available data provides the best estimate of phylogeny.The results show that Maesa is sister to all other taxa from the three families. Theophrastaceae are well supported, but both Myrsinaceae and Primulaceae are paraphyletic. We conclude that four families should be recognized, Maesaceae, Theophrastaceae, Primulaceae, and Myrsinaceae. For all families to be monophyletic, Samolus was transferred to Theophrastaceae, and Lysimachia, Anagallis, Trientalis, Glaux, Asterolinon, and Pelletiera were moved to the Myrsinaceae together with the genera Coris, Ardisiandra, and Cyclamen.

Journal Article↗

Phylogenetics of asterids based on 3 coding and 3 non-coding chloroplast DNA markers and the utility of non-coding DNA at higher taxonomic levels.

Asterids comprise 1/4-1/3 of all flowering plants and are classified in 10 orders and >100 families. The phylogeny of asterids is here explored with jackknife parsimony analysis of chloroplast DNA from 132 genera representing 103 families and all higher groups of asterids. Six different markers were used, three of the markers represent protein coding genes, rbcL, ndhF, and matK, and three other represent non-coding DNA; a region including trnL exons and the intron and intergenic spacers between trnT (UGU) to trnF (GAA); another region including trnV exons and intron, trnM and intergenic spacers between trnV (UAC) and atpE, and the rps16 intron. The three non-coding markers proved almost equally useful as the three coding genes in phylogenetic reconstruction at the high level of orders and families in asterids, and in relation to the number of aligned positions the non-coding markers were even more effective. Basal interrelationships among Cornales, Ericales, lamiids (new name replacing euasterids I), and campanulids (new name replacing euasterids II) are resolved with strong support. Family interrelationships are fully or almost fully resolved with medium to strong support in Cornales, Garryales, Gentianales, Solanales, Aquifoliales, Apiales, and Dipsacales. Within the three large orders Ericales, Lamiales, and Asterales, family interrelationships remain partly unclear. The analysis has contributed to reclassification of several families, e.g., Tetrameristaceae, Ebenaceae, Styracaceae, Montiniaceae, Orobanchaceae, and Scrophulariaceae (by inclusion of Pellicieraceae, Lissocarpaceae, Halesiaceae, Kaliphoraceae, Cyclocheilaceae, and Myoporaceae+Buddlejaceae, respectively), and to the placement of families that were unplaced in the APG-system, e.g., Sladeniaceae, Pentaphylacaceae, Plocospermataceae, Cardiopteridaceae, and Adoxaceae (in Ericales, Ericales, Lamiales, Aquifoliales, and Dipsacales, respectively), and Paracryphiaceae among campanulids. Several families of euasterids remain unclassified to order.

Chloroplasts↗

Polemoniaceae phylogeny and classification: implications of sequence data from the chloroplast gene ndhF.

The chloroplast gene ndhF was used to study phylogenetic relationships of the Polemoniaceae at two levels: among members of the Ericales and among genera of the family. Sequence data for interfamilial analyses consisted of 2266 bp for 14 members of the Ericales, including four species of the Polemoniaceae, plus three outgroup taxa. The Polemoniaceae were found to be related to Diospyros, Fouquieria, the Primulales, Rhododendron, and Impatiens, but relationships among taxa were generally not well supported. The precise position of the Polemoniaceae within the Ericales remains obscure. Data for intrafamilial analyses consisted of 1031 bp for 27 species of the Polemoniaceae, including at least one species from most genera of the family, plus five outgroup taxa. A single most parsimonious tree was identified. The analyses suggested that subfamily Cobaeoideae, excluding Loeselia, is monophyletic and that Huthia is sister to Cantua. Acanthogilia was sister to the remainder of subfamily Cobaeoideae. Subfamily Polemonioideae plus Loeselia formed four subclades that were strongly supported as monophyletic and represent the major lineages of the subfamily.

Journal Article↗

The phylogeny of the Asteridae sensu lato based on chloroplast ndhF gene sequences.

A phylogenetic study of Asteridae sensu lato was conducted based on chloroplast ndhF gene sequences for 116 ingroup and 13 outgroup species. Prior molecular studies based on rbcL sequences identified terminal groups corresponding to families, but were unable to resolve relationships among them. These results are largely consistent with earlier rbcL studies, but provide much greater resolution and stronger bootstrap support throughout the tree. The parsimony analysis found eight equally parsimonious trees, all of which recognize four major clades with the following relationship: (Cornales (Ericales (Euasterids I, Euasterids II))). Euasterids I includes (Garryales ((Solanales, Boraginaceae) (Gentianales, Lamiales))), although with weak support for relationships among the named clades. Euasterids II includes (Aquifoliales (Asterales (Apiales, Dipsacales))) with strong support for these relationships. Relationships within Ericales are weakly supported and merit further attention.

DNA, Chloroplast↗

Mechanisms of derived unitegmy among Impatiens species.

Morphological transitions associated with ovule diversification provide unique opportunities for studies of developmental evolution. Here, we investigate the underlying mechanisms of one such transition, reduction in integument number, which has occurred several times among diverse angiosperms. In particular, reduction in integument number occurred early in the history of the asterids, a large clade comprising approximately one-third of all flowering plants. Unlike the vast majority of other eudicots, nearly all asterids have a single integument, with the only exceptions in the Ericales, a sister group to the other asterids. Impatiens, a genus of the Ericales, includes species with one integument, two integuments, or an apparently intermediate bifid integument. A comparison of the development of representative Impatiens species and analysis of the expression patterns of putative orthologs of the Arabidopsis thaliana ovule development gene INNER NO OUTER (INO) has enabled us to propose a mechanism responsible for morphological transitions between integument types in this group. We attribute transitions between each of the three integument morphologies to congenital fusion via a combination of variation in the location of subdermal growth beneath primordia and the merging of primordia. Evidence of multiple transitions in integument morphology among Impatiens species suggests that control of underlying developmental programs is relatively plastic and that changes in a small number of genes may have been responsible for the transitions. Our expression data also indicate that the role of INO in the outgrowth and abaxial-adaxial polarity of the outer integument has been conserved between two divergent angiosperms, the rosid Arabidopsis and the asterid Impatiens.

Arabidopsis↗

Phylogenetic inference in Rafflesiales: the influence of rate heterogeneity and horizontal gene transfer.

BACKGROUND: The phylogenetic relationships among the holoparasites of Rafflesiales have remained enigmatic for over a century. Recent molecular phylogenetic studies using the mitochondrial matR gene placed Rafflesia, Rhizanthes and Sapria (Rafflesiaceae s. str.) in the angiosperm order Malpighiales and Mitrastema (Mitrastemonaceae) in Ericales. These phylogenetic studies did not, however, sample two additional groups traditionally classified within Rafflesiales (Apodantheaceae and Cytinaceae). Here we provide molecular phylogenetic evidence using DNA sequence data from mitochondrial and nuclear genes for representatives of all genera in Rafflesiales. RESULTS: Our analyses indicate that the phylogenetic affinities of the large-flowered clade and Mitrastema, ascertained using mitochondrial matR, are congruent with results from nuclear SSU rDNA when these data are analyzed using maximum likelihood and Bayesian methods. The relationship of Cytinaceae to Malvales was recovered in all analyses. Relationships between Apodanthaceae and photosynthetic angiosperms varied depending upon the data partition: Malvales (3-gene), Cucurbitales (matR) or Fabales (atp1). The latter incongruencies suggest that horizontal gene transfer (HGT) may be affecting the mitochondrial gene topologies. The lack of association between Mitrastema and Ericales using atp1 is suggestive of HGT, but greater sampling within eudicots is needed to test this hypothesis further. CONCLUSIONS: Rafflesiales are not monophyletic but composed of three or four independent lineages (families): Rafflesiaceae, Mitrastemonaceae, Apodanthaceae and Cytinaceae. Long-branch attraction appears to be misleading parsimony analyses of nuclear small-subunit rDNA data, but model-based methods (maximum likelihood and Bayesian analyses) recover a topology that is congruent with the mitochondrial matR gene tree, thus providing compelling evidence for organismal relationships. Horizontal gene transfer appears to be influencing only some taxa and some mitochondrial genes, thus indicating that the process is acting at the single gene (not whole genome) level.

Bayes Theorem↗

Fossil flowers of ericalean affinity from the Late Cretaceous of Southern Sweden.

Charcoalified fossil flowers of a new genus and species (Paradinandra suecica) with affinities to Ericales s.l. (sensu lato) are described from the Late Cretaceous (Santonian-Campanian) from Southern Sweden. The flowers are pentamerous, hypogynous, and actinomorphic. Aestivation of sepals and petals is imbricate-quincuncial. The androecium consists of an outer whorl with single episepalous stamens and an inner whorl with paired epipetalous stamens. Pollen is small and probably tricolpate. Three carpels form a syncarpous ovary with numerous campylotropous ovules on parietal placentae. The styles are free for most of their length. The structure of mature fruits and seeds is unknown. Clear distinction of sepals and petals, possible dehiscence of anthers by restricted slits, presence of a nectary, and the general floral construction (salverform corolla) with a canalized access to the floral center clearly indicate insect pollination of the fossil flowers. Comparisons with extant taxa demonstrate that Paradinandra suecica shares many similarities with Ericales s.l. and in particular with members of Ternstroemiaceae, Theaceae, and Actinidiaceae. However, it is neither identical to any one genus of these families nor to any of the previously described ericalean taxa from the Cretaceous and thus provides further evidence of the diversity of Cretaceous ericalean plants.

Journal Article↗

Co-occurrence of three fungal root symbionts in Gaultheria poeppiggi DC in central Argentina.

The roots of Gaultheria poeppiggi (Ericaceae) were examined for fungal symbiont colonization. Typical structures of ericoid mycorrhizas (hyphae and intracellular coil hyphae complexes), dark septate fungal endophytes (hyphae and sclerotia), and arbuscular mycorrhizas (hyphae, coils, vesicles and arbuscules) were found in the roots of all the individuals examined. The evolutionarily derived position of Gaultheria within the Ericales may suggest that G. poeppiggi recently acquired the ability to form arbuscular mycorrhizas rather than having retained it from ancestral lines.

Argentina↗

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↗

Purification and characterization of phytocystatins from kiwifruit cortex and seeds.

Kiwifruit cysteine proteinase inhibitors (KCPIs) were purified from the cortex and seeds of kiwifruit after inactivation of the abundant cortex cysteine proteinase actinidain. One major (KCPI1) and four minor cystatins were identified from Actinidia deliciosa ripe mature kiwifruit cortex as well as a seed KCPI from A. chinensis. The predominant cortex cystatin, KCPI1, inhibited clan CA, family C1 (papain family) cysteine proteinases (papain, chymopapain, bromelain, ficin, human cathepsins B, H and L, actinidain and the house dust mite endopeptidase 1), while cysteine proteinases belonging to other families, [clostripain (C11), streptopain (C10) and calpain (C2)] were not inhibited. Inhibition constants (K(I)) ranged between 0.001 nM for cathepsin L and 0.98 nM for endopeptidase 1. The K(I) (14 nM) for KCPI1 inhibiting actinidain is at least 2 orders of magnitude higher than for other plant proteinases measured. The cortex KCPI1 and a seed KCPI purified from seeds had the same N-terminal sequence (VAAGGWRPIESLNSAEVQDV). BLAST-matching the peptide sequence against an in-house generated Actinidia EST database, identified 81 cDNAs that exactly matched the measured KCPI1 peptide sequence. Peptide sequences of two other cortex KCPIs each exactly matched a predicted peptide sequence of a cDNA from kiwifruit. The predicted peptide sequence of KCPI1 of 116 amino acids encodes a signal peptide and does not contain cysteine. Without the signal peptide (mature protein), KCPI1 has a molecular mass of approximately 11 kDa, possesses the consensus sequence characteristic for the phytocystatins and shows the highest homology to a cystatin from Citrusxparadisi (52% identity). This is the first report of phytocystatins from the Ericales.

Actinidia↗