We report the draft genome of the black cottonwood tree, Populus trichocarpa . Integration of shotgun sequence assembly with genetic mapping enabled chromosome-scale reconstruction of the genome. More than 45,000 putative protein-coding genes were identified. Analysis of the assembled genome revealed a whole-genome duplication event; about 8000 pairs of duplicated genes from that event survived in the Populus genome. A second, older duplication event is indistinguishably coincident with the divergence of the Populus and Arabidopsis lineages. Nucleotide substitution, tandem gene duplication, and gross chromosomal rearrangement appear to proceed substantially more slowly in Populus than in Arabidopsis. Populus has more protein-coding genes than Arabidopsis , ranging on average from 1.4 to 1.6 putative Populus homologs for each Arabidopsis gene. However, the relative frequency of protein domains in the two genomes is similar. Overrepresented exceptions in Populus include genes associated with lignocellulosic wall biosynthesis, meristem development, disease resistance, and metabolite transport.
A r t i c l e s Theobroma cacao L. is a diploid tree fruit species (2n = 2x = 20 (ref. 1)) endemic to the South American rainforests. Cocoa was domesticated approximately 3,000 years ago 2 in Central America 3. The Criollo cocoa variety, having a nearly unique and homozygous genotype, was among the first to be cultivated 4. Criollo is now one of the two cocoa varieties providing fine flavor chocolate. However, due to its poor agronomic performance and disease susceptibility, more vigorous hybrids created with foreign (Forastero) genotypes have been introduced. These hybrids, named Trinitario, are now widely cultivated 5. Here we report the sequence of a Belizean Criollo plant 6. Consumers have shown an increased interest for high-quality chocolate, and for dark chocolate, containing a higher percentage of cocoa 7. Fine-cocoa production is nevertheless estimated to be less than 5% of the world cocoa production due to the low productivity and disease susceptibility of the traditional fine-flavor cocoa varieties. Therefore, breeding of improved Criollo varieties is important for sustainable production of fine-flavor cocoa. 3.7 million tons of cocoa are produced annually (see URLs). However, fungal, oomycete and viral diseases, as well as insect pests, are responsible for an estimated 30% of harvest losses (see URLs). Like many other tropical crops, knowledge of T. cacao genetics and genomics is limited. To accelerate progress in cocoa breeding and the understanding of its biochemistry, we sequenced and analyzed the genome
Genomic comparisons provide evidence for ancient genome-wide duplications in a diverse array of animals and plants. We developed a birth-death model to identify evidence for genome duplication in EST data, and applied a mixture model to estimate the age distribution of paralogous pairs identified in EST sets for species representing the basal-most extant flowering plant lineages. We found evidence for episodes of ancient genome-wide duplications in the basal angiosperm lineages including Nuphar advena (yellow water lily: Nymphaeaceae) and the magnoliids Persea americana (avocado: Lauraceae), Liriodendron tulipifera (tulip poplar: Magnoliaceae), and Saruma henryi (Aristolochiaceae). In addition, we detected independent genome duplications in the basal eudicot Eschscholzia californica (California poppy: Papaveraceae) and the basal monocot Acorus americanus (Acoraceae), both of which were distinct from duplications documented for ancestral grass (Poaceae) and core eudicot lineages. Among gymnosperms, we found equivocal evidence for ancient polyploidy in Welwitschia mirabilis (Gnetales) and no evidence for polyploidy in pine, although gymnosperms generally have much larger genomes than the angiosperms investigated. Cross-species sequence divergence estimates suggest that synonymous substitution rates in the basal angiosperms are less than half those previously reported for core eudicots and members of Poaceae. These lower substitution rates permit inference of older duplication events. We hypothesize that evidence of an ancient duplication observed in the Nuphar data may represent a genome duplication in the common ancestor of all or most extant angiosperms, except Amborella.
Annual plants grow vegetatively at early developmental stages and then transition to the reproductive stage, followed by senescence in the same year. In contrast, after successive years of vegetative growth at early ages, woody perennial shoot meristems begin repeated transitions between vegetative and reproductive growth at sexual maturity. However, it is unknown how these repeated transitions occur without a developmental conflict between vegetative and reproductive growth. We report that functionally diverged paralogs FLOWERING LOCUS T1 (FT1) and FLOWERING LOCUS T2 (FT2), products of whole-genome duplication and homologs of Arabidopsis thaliana gene FLOWERING LOCUS T (FT), coordinate the repeated cycles of vegetative and reproductive growth in woody perennial poplar (Populus spp.). Our manipulative physiological and genetic experiments coupled with field studies, expression profiling, and network analysis reveal that reproductive onset is determined by FT1 in response to winter temperatures, whereas vegetative growth and inhibition of bud set are promoted by FT2 in response to warm temperatures and long days in the growing season. The basis for functional differentiation between FT1 and FT2 appears to be expression pattern shifts, changes in proteins, and divergence in gene regulatory networks. Thus, temporal separation of reproductive onset and vegetative growth into different seasons via FT1 and FT2 provides seasonality and demonstrates the evolution of a complex perennial adaptive trait after genome duplication.ife cycles of higher plants display a great diversity in morphological and seasonal adaptation. Annual plants grow, reproduce, and senesce within a growing season, whereas woody perennials display successive years of vegetative growth before reaching sexual maturity (1-3). After this time, shoot meristems begin cyclical transitions between vegetative and reproductive growth. Consequently, shoots may repeatedly form early vegetative buds (Vegetative Zone I), reproductive buds (Floral Zone), and late vegetative buds (Vegetative Zone II) in a sequential manner (3). However, our understanding of the mechanisms underlying such complex phenotypes, and thus variation in growth habits and adaptation, remain rudimentary. In the herbaceous perennial Arabis alpina, repeated transcriptional repression and activation of PERPETUAL FLOWERING 1 (PEP1), an ortholog of the floral repressor FLOWERING LOCUS C (FLC) in annual Arabidopsis thaliana (4), controls recurring seasonal transitions between reproductive and vegetative phases (5). However, a true functional ortholog of FLC has not been reported in trees, nor does phylogenetic analysis point to a clear structural ortholog of FLC in poplar (Populus spp.) (6).Previous results showed that FLOWERING LOCUS T1 (FT1) (7) and FLOWERING LOCUS T2 (FT2) (8) under the cauliflower mosaic virus 35S (CaMV 35S) constitutive overexpression promoter induce early flowering in poplar. Transcript abundance of both genes gradually increases in the growing season as poplar trees mature....
Plant volatiles play important roles in signalling between plants and insects, but their role in communication among plants remains controversial. Previous research on plant-plant communication has focused on interactions between neighbouring plants, largely overlooking the possibility that volatiles function as signals within plants. Here, we show that volatiles released by herbivore-wounded leaves of hybrid poplar (Populus deltoides x nigra) prime defences in adjacent leaves with little or no vascular connection to the wounded leaves. Undamaged leaves exposed to volatiles from wounded leaves on the same stem had elevated defensive responses to feeding by gypsy moth larvae (Lymantria dispar L.) compared with leaves that did not receive volatiles. Volatile signals may facilitate systemic responses to localized herbivory even when the transmission of internal signals is constrained by vascular connectivity. Self-signalling via volatiles is consistent with the short distances over which plant response to airborne cues has been observed to occur and has apparent benefits for emitting plants, suggesting that within-plant signalling may have equal or greater ecological significance than signalling between plants.
Summary• Herbivore-induced plant volatiles (HIPVs), in addition to attracting natural enemies of herbivores, can serve a signaling function within plants to induce or prime defenses. However, it is largely unknown, particularly in woody plants, which volatile compounds within HIPV blends can act as signaling molecules.• Leaves of hybrid poplar saplings were exposed in vivo to naturally wound-emitted concentrations of the green leaf volatile (GLV) cis-3-hexenyl acetate (z3HAC) and then subsequently fed upon by gypsy moth larvae. Volatiles were collected throughout the experiments, and leaf tissue was collected to measure phytohormone concentrations and expression of defense-related genes.• Relative to controls, z3HAC-exposed leaves had higher concentrations of jasmonic acid and linolenic acid following gypsy moth feeding. Furthermore, z3HAC primed transcripts of genes that mediate oxylipin signaling and direct defenses, as determined by both qRT-PCR and microarray analysis using the AspenDB 7 K expressed sequence tags (EST) microarray containing c. 5400 unique gene models. Moreover, z3HAC primed the release of terpene volatiles.• The widespread priming response suggests an adaptive benefit to detecting z3HAC as a wound signal. Thus, woody plants can detect and use z3HAC as a signal to prime defenses before actually experiencing damage. GLVs may therefore have important ecological functions in arboreal ecosystems.
The aromatic polymer lignin protects plants from most forms of microbial attack. Despite the fact that a significant fraction of all lignocellulose degraded passes through arthropod guts, the fate of lignin in these systems is not known. Using tetramethylammonium hydroxide thermochemolysis, we show lignin degradation by two insect species, the Asian longhorned beetle (Anoplophora glabripennis) and the Pacific dampwood termite (Zootermopsis angusticollis). In both the beetle and termite, significant levels of propyl side-chain oxidation (depolymerization) and demethylation of ring methoxyl groups is detected; for the termite, ring hydroxylation is also observed. In addition, culture-independent fungal gut community analysis of A. glabripennis identified a single species of fungus in the Fusarium solani/Nectria haematococca species complex. This is a soft-rot fungus that may be contributing to wood degradation. These results transform our understanding of lignin degradation by wood-feeding insects.Asian longhorned beetle ͉ Pacific dampwood termite ͉ TMAH thermochemolysis ͉ Anoplophora glabripennis ͉ Zootermopsis angusticollis L ignin plays a central role in carbon cycling on Earth. Its heterogeneous structure imparts plants with structural rigidity and also serves to protect cellulose and hemicellulose from degradation (1). Most of what is known about lignin biodegradation is from pure culture studies with filamentous basidiomycete fungi, known as white-rot and brown-rot decay. Although both white-rot and brown-rot fungal degradation have been characterized, much more is known about the white-rot system (2, 3). White-rot fungi simultaneously degrade the three major components of the plant cell wall: lignin, cellulose, and hemicellulose. Analysis of white-rot-degraded wood shows that the reactions in lignin: (i) are oxidative, (ii) involve demethylation (or demethoxylation), (iii) include side-chain oxidation at C ␣ , and (iv) involve propyl side-chain cleavage between C ␣ and C  (Fig. 1) (4). In contrast to white-rot fungi, brown-rot fungi are able to circumvent the lignin barrier, removing the hemicellulose and cellulose with only minor modification to the lignin. Consequently, lignin remains a major component of the degraded plant cell wall (5). The remaining lignin is demethylated on aryl methoxy groups and contains a greater number of ring hydroxyl groups (6).Little is known about lignin degradation in complex ecosystems, such as insect guts, where a consortium of microbes may be involved in degradation rather than just a single species. Although cellulose degradation in insect guts is well documented (7,8), the fate of lignin has not clearly been demonstrated (9, 10), and it is widely accepted that insect gut systems do not have the capacity to degrade lignin (10). Although the majority of previous reports suggest that many wood-feeding insects overcome the lignin barrier by feeding on predegraded wood (11) or through exosymbiotic relationships with wood-degrading fungi (12, 13), there are species of insects...
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