Biology subjects

Zhang, S.

Publications and source records attributed to Zhang, S..

At least 415 records · Page 23Linked to original sources

bioRxiv2025

Dynamic μ-PBWT: Dynamic Run-length Compressed PBWT for Biobank Scale Data

Durbins positional Burrows-Wheeler transform (PBWT) supports efficient haplotype matching and queries given a panel of haplotypes. It has been widely used for statistical phasing, imputation and identity-by-descent (IBD) detection. However, the original PBWT panel doesnt support dynamic updates when haplotypes need to be added or deleted from the panel. Dynamic-PBWT (d-PBWT) solved this problem but it is not memory efficient. While the memory constraint problem of the PBWT has been tackled by Syllable-PBWT and {micro}-PBWT, these are static data structures that do not allow updates. Additionally, Syllable-PBWT only supports long-match query and {micro}-PBWT only supports set-maximal match query, limiting their functionality in the compressed form. In this paper, we present Dynamic {micro}-PBWT (which can also be seen as compressed d-PBWT) that is memory efficient and supports dynamic updates. We run-length compress PBWT to achieve better compression rate and store the runs in the self-balancing trees to enable dynamic updates. We show that the number of updates per insertion or deletion in the tree at each site is constant regardless of the number of haplotypes in the panel and the updates can be made without decompressing the index. In addition, we use orders of magnitude less memory than d-PBWT. We also provide a long match query algorithm that can easily be extended back to the original {micro}-PBWT. Overall, the flexibility and space-efficiency of Dynamic {micro}-PBWT makes it a potential index data structure for biobank scale genetic data analyses. The source code for Dynamic {micro}-PBWT is available at https://github.com/ucfcbb/Dynamic-mu-PBWT.

bioinformatics↗

bioRxiv2025

Recent thymic emigrants are preferentially recruited into the memory pool during persistent infection

Cytomegalovirus (CMV) leads to a unique phenomenon known as memory inflation, where antigen-specific memory CD8+ T cells continue to accumulate in the peripheral tissues during the latent stage of infection. However, it is still not clear how the inflating pool of memory CD8+ T cells is generated and maintained. In this study, we used murine cytomegalovirus (MCMV) as a model of persistent infection and fate-mapping mice to determine the dynamics of CD8+ T cell recruitment into the memory pool. We found that neonatal exposure to CMV leads to an expansion of newly made CD8+ T cells (recent thymic emigrants, RTEs), which are maintained in the long-lived memory compartment. In contrast, CD8+ T cells made during the latent phase of infection (mature CD8+ T cells) contribute little to the memory pool. We also observed notable phenotypic differences between RTEs and mature cells. Whereas RTEs present at the time of infection gave rise to more effector memory cells, the cells produced later in infection were biased towards becoming central memory cells. Importantly, the preferential recruitment of RTEs into the effector memory pool also occurs during adult exposure to CMV. Collectively, these data demonstrate that persistent infection expands the RTE population, and timing of infection dictates whether neonatal or adult RTEs are locked in to the memory pool. Author SummaryFollowing infection with CMV, CD8+ T cells accumulate in the blood and peripheral organs over time, a feature termed memory inflation. However, it is not clear whether memory inflation is due to the continuous recruitment of cells made during the latent stage of infection or expansion of CD8+ T cells that were present at the time of infection. To address this question, we used a fate-mapping mouse model and examined the recruitment of CD8+ T cells that were produced during different stages of infection. Surprisingly, we discovered that CD8+ T cells exported from the thymus just prior to infection are preferentially recruited and maintained in the memory pool. In contrast, CD8+ T cells made during the latent stage contribute minimally to the inflating pool and exhibit a less differentiated phenotype. These results provide a new conceptual framework for understanding how the memory pool is generated and maintained after persistent viral infection.

immunology↗

bioRxiv2025

Haplotype Matching with GBWT for Pangenome Graphs

Traditionally, variations from a linear reference genome were used to represent large sets of haplotypes compactly. In the linear reference genome based paradigm, the positional Burrows-Wheeler transform (PBWT) has traditionally been used to perform efficient haplotype matching. Pangenome graphs have recently been proposed as an alternative to linear reference genomes for representing the full spectrum of variations in the human genome. However, haplotype matches in pangenome graph based haplotype sets are not trivially generalizable from haplotype matches in the linear reference genome based haplotype sets. Work has been done to represent large sets of haplotypes as paths through a pangenome graph. The graph Burrows-Wheeler transform (GBWT) is one such work. The GBWT essentially stores the haplotype paths in a run length compressed BWT with compressed local alphabets. Although efficient in practice count and locate queries on the GBWT were provided by the original authors, the efficient haplotype matching capabilities of the PBWT have never been shown on the GBWT. In this paper, we formally define the notion of haplotype matches in pangenome graph-based haplotype sets by generalizing from haplotype matches in linear reference genome-based haplotype sets. We also describe the relationship between set maximal matches, long matches, locally maximal matches, and text maximal matches on the GBWT, PBWT, and the BWT. We provide algorithms for outputting some of these matches by applying the data structures of the r-index (introduced by Gagie et al.) to the GBWT. We show that these structures enable set maximal match and long match queries on the GBWT in almost linear time and in space close to linear in the number of runs in the GBWT. We also provide multiple versions of the query algorithms for different combinations of the available data structures. The long match query algorithms presented here even run on the BWT in the same time complexity as the GBWT due to their similarity.

bioinformatics↗

bioRxiv2025

Reconstructing the three-dimensional architecture of extrachromosomal DNA with ec3D

Extrachromosomal DNAs (ecDNAs) are large, acentric, circular DNA molecules that occur pervasively across many human cancers. EcDNA can drive tumor formation and evolution, contribute to drug resistance, and associate with poor patient survival outcomes. Beyond mediating high copy numbers, the circular topology and dynamic conformational changes of ecDNA disrupt topological domains and rewire regulatory networks, thereby conferring an important role in the transcriptional regulation of oncogenes. Here, we develop ec3D, a computational method for reconstructing the three-dimensional structures of ecDNA and analyzing significant interactions from high-throughput chromatin capture (Hi-C) data. Given a candidate ecDNA sequence and the corresponding whole-genome Hi-C as input, ec3D reconstructs the spatial structure of ecDNA by maximizing the Poisson likelihood of observed interactions. Ec3Ds performance was validated using both simulated ecDNA structures with varying conformations and Hi-C data from previously-characterized cancer cell lines. Our reconstructions reveal that ecDNAs occupy spherical configurations and mediate unique long-range interactions involved in gene regulation. Through algorithmic innovations, ec3D can resolve complex ecDNA structures with duplicated genomic segments, identify multi-way interactions, and distinguish between interactions arising from direct spatial proximity and secondary interactions resulting from alternative folding patterns or intermolecular (trans) contacts of ecDNA molecules. Our findings provide insights into how the spatial organization of ecDNA may influence gene regulation and contribute to increased oncogene expression. Code availabilityhttps://github.com/AmpliconSuite/ec3D

cancer biology↗

bioRxiv2025

Epithelial miR-149-5p up-regulation is associated with immune evasion in progressive bronchial premalignant lesions

The molecular drivers bronchial premalignant lesion progression to invasive lung squamous cell carcinoma are not well defined. Prior work profiling longitudinally collected bronchial premalignant lesion biopsies by RNA sequencing defined a proliferative subtype, enriched with bronchial dysplasia. We found that a gene co-expression module associated with interferon gamma signaling and antigen processing/presentation was down-regulated in progressive/persistent versus regressive lesions within the proliferative subtype, suggesting a functional impact of these genes on immune evasion. RNA from these same premalignant lesions was profiled by microRNA (miRNA) sequencing and a miRNA-gene network analysis identified hsa-miR-149-5p as a potential regulator of this antigen presentation gene co-expression module associated with lesion progression. hsa-miR-149-5p was found to be predominantly expressed in the epithelium and up-regulated in progressive/persistent versus regressive proliferative lesions while targets of this miRNA, the transcriptional coactivator of MHC-I gene expression, NLRC5, and the genes it regulates were down-regulated. MicroRNA in situ hybridization of hsa-miR-149-5p in tissue from adjacent fixed biopsies showed that hsa-miR-149-5p was increased in areas of bronchial dysplasia in progressive/persistent versus regressive lesions. Imaging mass cytometry showed that NLRC5 protein expression was decreased in progressive/persistent versus regressive lesions within areas of hyperplasia, metaplasia, and dysplasia. Additionally, basal cells with high versus low levels of NLRC5 were found to be in close spatial proximity to CD8 T cells, suggesting that these cells exhibit increased functional MHC-I gene expression in lesions with low hsa-miR-149-5p expression. Collectively, our data suggests a functional role for hsa-miR-149-5p in bronchial premalignant lesions and may serve as a therapeutic target for PML immunomodulation. STATEMENT OF SIGNIFICANCEIntegrative analysis across bronchial premalignant lesions has identified and localized a potential regulator of immune evasion in progressive/persistent lesions that could be a novel therapeutic target.

cancer biology↗

bioRxiv2025

sciLaMA: A Single-Cell Representation Learning Framework to Leverage Prior Knowledge from Large Language Models

Single-cell RNA sequencing (scRNA-seq) enables high-resolution exploration of cellular diversity and gene regulation, yet analyzing such data remains challenging due to technical and methodological limitations. Existing task-specific deep generative models like Variational Auto-Encoder (VAE) and its variants struggle to incorporate external biological knowledge, while transformer-based foundational large Language Models (LLMs or large LaMs) face limitations in computational cost and applicability to tabular gene expression data. Here, we introduce sciL-aMA (single-cell interpretable Language Model Adapter), a novel representation learning frame-work that bridges these gaps by integrating static gene embeddings from multimodal LLMs with scRNA-seq tabular data through a paired-VAE architecture. Our approach generates context-aware representations for both cells and genes and outperforms state-of-the-art methods in key single-cell downstream tasks, including batch effect correction, cell clustering, and cell-state-specific gene marker and module identification, while maintaining computational efficiency. sciL-aMA offers a computationally efficient, unified framework for comprehensive single-cell data analysis and biologically interpretable gene module discovery. Source code is available at https://github.com/microsoft/sciLaMA

bioinformatics↗

bioRxiv2025

Differences between Scaly-sided Merganser (Mergus squamatus) and Common Merganser (M. merganser) feather microstructure

The microstructures of breast feathers, flank feathers, rectrices, and down feathers of both Scaly-sided Merganser (Mergus squamatus) and Common Merganser (M. merganser) were meticulously examined using scanning electron microscope. The observed data were subjected to the Mann-Whitney U test, followed by hierarchical clustering and Fishers discriminant analysis of indicators with significant differences in morphological structure. The findings revealed statistically significant differences between Scaly-sided Merganser and Common Merganser in terms of the length of the base handle from distal barbules in the breast feathers, the length of the base handle, the number of hooked barbules, the number of cilia from distal barbules, and the number of ventral teeth from proximal barbules in rectrices. Additionally, variations in the distance between nodules, the nodular diameter, and the length of the base nodular of the down feathers were noted. Hierarchical clustering successfully categorized Scaly-sided Merganser and Common Merganser into distinct groups, achieving a 100% accuracy rate. Moreover, Fishers discriminant analysis demonstrated a 100% correctness rate when the back-substitution method was used to test the discriminant equation. Consequently, the length of the base handle of distal barbules in breast feathers, along with all selected indicators from rectrices and down feathers, may provide references to identify Scaly-sided Merganser and Common Merganser.

zoology↗

bioRxiv2025

Structural basis of RECQL5-induced RNA polymerase II transcription braking and subsequent reactivation

During productive transcription elongation, the speed of RNA polymerase II (Pol II) is highly dynamic within individual genes and varies between different genes1,2. Unregulated rapid transcription elongation can lead to detrimental consequences such as transcription-replication collisions, altered alternative splicing patterns, and genome instability1-7. Therefore, elongating Pol II requires mechanisms to slow its progression, yet the molecular basis of transcription braking remains unclear. RECQL5 is a DNA helicase that functions as a general elongation factor by slowing down Pol II8-11. Here we report cryo-electron microscopy (cryo-EM) structures of human RECQL5 bound to multiple transcription elongation complexes. Combined with biochemical analysis, we identify an -helix of RECQL5 responsible for Pol II binding and slowdown of transcription elongation. We further reveal that the transcription-coupled DNA repair (TCR) complex allows Pol II to overcome RECQL5-induced transcription braking through concerted actions of its translocase activity and competition with RECQL5 for engaging Pol II. Additionally, RECQL5 inhibits TCR-mediated Pol II ubiquitination to prevent activation of the DNA repair pathway. Our results suggest a model in which RECQL5 and the TCR complex coordinately regulate the transcription elongation rate to ensure transcription efficiency while maintaining genome stability. This work provides a framework for future studies on the regulatory role of elongation speed in gene expression.

molecular biology↗

bioRxiv2025

Multi-species integration, alignment and annotation of single-cell RNA-seq data with CAMEX

Single-cell RNA-seq (scRNA-seq) data from multiple species present remarkable opportunities to explore cellular origins and evolution. However, integrating and annotating scRNA-seq data across different species remains challenging due to the variations in sequencing techniques, ambiguity of homologous relationships, and limited biological knowledge. To tackle the above challenges, we introduce CAMEX, a heterogeneous Graph Neural Network (GNN) tool that leverages many-to-many homologous relationships for multi-species integration, alignment, and annotation of scRNA-seq data from multiple species. Notably, CAMEX outperforms state-of-the-art methods integration on various cross-species benchmarking datasets (ranging from one to eleven species). Besides, CAMEX facilitates the alignment of diverse species across different developmental stages, significantly enhancing our understanding of organ and organism origins. Furthermore, CAMEX enables the detection of species-specific cell types and marker genes through cell and gene embedding. In short, CAMEX holds the potential to provide invaluable insights into how evolutionary forces operate across different species at single-cell resolution.

bioinformatics↗

bioRxiv2025

Dietary protein source mediates colitis pathogenesis through bacterial modulation of bile acids

Introductory paragraphEvidence-based dietary recommendations for individuals with inflammatory bowel diseases (IBD) are limited. Red meat consumption is associated with increased IBD incidence and relapse in patients, suggesting that switching to a plant-based diet may limit gut inflammation. However, the mechanisms underlying the differential effects of these diets remain poorly understood. Feeding diets containing plant- or animal-derived proteins to murine colitis models revealed that mice given a beef protein (BP) diet exhibited the most severe colitis, while mice fed pea protein (PP) developed mild inflammation. The colitis-promoting effects of BP were microbially-mediated as determined by bacterial elimination or depletion and microbiota transplant studies. In the absence of colitis, BP-feeding reduced abundance of Lactobacillus johnsonii and Turicibacter sanguinis and expanded Akkermansia muciniphila, which localized to the mucus in association with decreased mucus thickness and quality. BP-fed mice had elevated primary and conjugated fecal bile acids (BAs), and taurocholic acid administration to PP-fed mice worsened colitis. Dietary psyllium protected against BP-mediated inflammation, restored BA-modulating commensals and normalized BA ratios. Collectively, these data suggest that the protein component of red meat may be responsible, in part, for the colitis-promoting effects of this food source and provide insight into dietary factors that may influence IBD severity.

microbiology↗

bioRxiv2025

Sequence-based Drug-Target Complex Pre-training Enhances Protein-Ligand Binding Process Predictions Tackling Crypticity

Predicting protein-ligand binding characteristics, such as affinity and kinetics, is critical for accelerating drug discovery. However, many existing computational methods face key limitations, including insufficient integration of comprehensive databases, inadequate representation of protein structural dynamics, and incomplete modeling of microscale protein-ligand interactions. To address these challenges, we introduce ProMoNet, a sequence-based pre-training and fine-tuning framework to enhance the prediction of protein-ligand binding characteristics. ProMoNet leverages protein and molecular foundation models to expand data coverage and enhance diversity. It also introduces a pre-training strategy based on protein-ligand binding site prediction, which bridges protein- and ligand-level representations to support downstream prediction tasks involving protein-ligand complexes. Our pre-training module effectively models microscale protein-ligand interactions and captures the dynamic nature of proteins, including binding site crypticity, without relying on 3-dimensional structural inputs. Notably, this module surpasses or matches state-of-the-art structure-based methods in identifying exposed and cryptic binding sites while maintaining high efficiency. Our fine-tuning module then efficiently transfers the pre-trained knowledge to downstream tasks such as binding affinity and binding kinetics prediction, achieving superior performance. The combination of ProMoNets strong performance and demonstrated efficiency across multiple tasks highlights its potential for broad applications in drug discovery. Scientific ContributionWe propose ProMoNet, a sequence-based pre-training and fine-tuning framework for protein-ligand binding characteristic prediction, where protein-ligand binding site prediction is introduced as a pre-training strategy to bridge independent protein- and ligand-level representations for downstream complex-level tasks. We design two dedicated modules, including a pre-training module that models microscale protein-ligand interactions and captures protein dynamics, as well as a fine-tuning module that efficiently integrates the pre-trained representations for downstream tasks. Even compared to structure-based methods, ProMoNet matches state-of-the-art performance in exposed and cryptic binding site identification and delivers superior results in binding affinity and kinetics prediction, making it a promising tool for drug discovery.

bioinformatics↗

bioRxiv2025

5' UTR-mediated retention of eIF3 on 80S ribosomes promotes co-translational folding of ER membrane proteins

Co-translational folding of nascent polypeptides is essential for protein function and cellular homeostasis. Ribosome-associated chaperones assist in this process, but coordination between their recruitment and translation initiation remains poorly understood. We report here that specific binding sites for eukaryotic translation initiation factor eIF3 within the 5 untranslated regions (5UTRs) of mRNAs promote its retention on 80S ribosomes during the synthesis of select endoplasmic reticulum (ER) membrane proteins. Disruption of these eIF3 binding sites leads to misfolding and sequestration of newly synthesized membrane proteins into ER whorls. Sequestration into ER whorls is exacerbated by HSP70 inhibition but can be rescued by overexpressing HSPA8 and HSPA1. Cross-linking assays reveal that 5UTR binding sites stabilize eIF3-80S interactions during early elongation, facilitating recruitment of HSPA8 to ribosomes. These findings indicate that genetic instructions within 5UTRs direct eIF3-mediated chaperone recruitment, ensuring proper co-translational folding of ER membrane proteins.

cell biology↗

bioRxiv2025

A Preliminary Application of Bird Surveys Using Environmental DNA Technology: in Futian Mangrove Nature Reserve

Futian Mangrove Nature Reserve is a vital habitat for birds in Shenzhen city of China, and the diversity of bird species is of considerable concern. Traditional surveys depend on direct observation and expert assessment, necessitating significant manpower for tracking. This study utilized environmental DNA metabarcoding technology to gather 128 environmental samples of water, soil, and air, along with samples of bird feces and feathers collected during the summer and winter of 2023. A total of 72 bird operational taxonomic units (OTUs) were identified. The results revealed variations in bird detection across different environmental samples, with air samples yielding 52 bird OTUs, water samples 29, and soil samples 23. Detected bird OTUs were notably higher in winter samples compared to summer. Additionally, the average number of bird OTUs detected in air samples during summer and winter was 14.9 and 10.1, respectively, significantly exceeding those found in the corresponding water and soil samples. The identification of bird feces and feather samples confirmed the presence of the identified bird species in the environmental samples. This study concludes that environmental DNA technology can serve as a valuable complementary method for monitoring bird diversity, with different environmental samples showing distinct preferences in bird detection.

ecology↗

bioRxiv2025

Detection of alpha synuclein seeding activity in tear fluid in patients with Parkinson's disease

Detection of alpha-synuclein seeding activity in tear fluid (TF) might provide a promising non-invasive biomarker for Parkinsons disease (PD) diagnosis. In this study, we applied the alpha-synuclein seeding amplification assay (aSynSAA) to detect misfolded alpha-synuclein (aSyn) aggregation in TF from PD patients. The discovery cohort included 11 PD patients and 13 controls, and the validation cohort consisted of 9 PD patients and 11 controls without synucleinopathies. The aSynSAA yielded positive results in over 55% of PD patients. These findings were confirmed in a second cohort, including patients with prion diseases as a negative control for synuclein pathology. Our results demonstrate for the first time the ability of aSynSAA to distinguish between PD and control groups in TF, with PD showing the highest seeding activity compared to prion disease and control groups. Further comparisons between cerebrospinal fluid (CSF) and TF samples from the same individuals revealed consistent seeding results across both biofluids. These findings highlight the potential of tear fluid as a novel, accessible medium for detecting Lewy body-specific misfolded synuclein aggregation in PD, which could aid in early diagnosis and disease progression monitoring.

neuroscience↗

bioRxiv2025

Toward highly accurate reference and method selection for universal cross-dataset cell type annotation

Cell type annotation is a critical and essential task in single-cell data analysis. Various reference-based methods have provided rapid annotation for diverse single-cell data. However, how to select the optimal references and methods is often overlooked. To this end, we present a cross-dataset cell-type annotation methodology with a universal reference data and method selection strategy (CAMUS) to achieve highly accurate and efficient annotations. We demonstrate the advantages of CAMUS by conducting comprehensive analyses on 672 pairs of cross-species scRNA-seq datasets. The annotation results with references selected by CAMUS achieved substantial accuracy gains (25.0-124.7%) over random selection strategies across five reference-based methods. CAMUS achieved high accuracy in choosing the best reference-method pair among 3360 pairs (49.1%). Moreover, CAMUS showed high accuracy in selecting the best methods on the 80 scST datasets (82.5%) and five scATAC-seq datasets (100.0%), illustrating its universal applicability. In addition, we utilized the CAMUS score with other metrics to predict the annotation accuracy, providing direct guidance on whether to accept current annotation results.

bioinformatics↗

bioRxiv2025

A high-quality reference genome and comparative genomics of the widely-farmed bandedcricket (Gryllodes sigillatus) identifies selective breeding targets

Farmed insects have gained attention as an alternative, sustainable source of protein with a lower carbon footprint than traditional livestock. We present a high-quality reference genome for one of the most commonly farmed insects, the banded cricket Gryllodes sigillatus. In addition to its agricultural importance, G. sigillatus is also a model in behavioural and evolutionary ecology research on reproduction and mating systems. We report comparative genomic analyses that clarify the banded crickets evolutionary history, identify gene family expansions and contractions unique to this lineage, associate these with agriculturally important traits, and identify targets for genome-assisted breeding efforts. The high-quality G. sigillatus genome assembly plus accompanying comparative genomic analyses serve as foundational resources for both applied and basic research on insect farming and behavioural biology, enabling researchers to pinpoint trait-associated genetic variants, unravel functional pathways governing those phenotypes, and accelerate selective breeding efforts to increase the efficacy of large-scale insect farming operations.

genomics↗

bioRxiv2025

Dose-dependent sensitivity of human 3D chromatin to a heart disease-linked transcription factor

Dosage-sensitive transcription factors (TFs) underlie altered gene regulation in human developmental disorders, and cell-type specific gene regulation is linked to the reorganization of 3D chromatin during cellular differentiation. Here, we show dose-dependent regulation of chromatin organization by the congenital heart disease (CHD)- linked, lineage-restricted TF TBX5 in human cardiomyocyte differentiation. Genome organization, including compartments, topologically associated domains, and chromatin loops, are sensitive to reduced TBX5 dosage in a human model of CHD, with variations in response across individual cells. Regions normally bound by TBX5 are especially sensitive, while co-occupancy with CTCF partially protects TBX5-bound TAD boundaries and loop anchors. These results highlight the importance of lineage-restricted TF dosage in cell-type specific 3D chromatin dynamics, suggesting a new mechanism for TF-dependent disease.

genomics↗

bioRxiv2025

Ustilaginoidea virens suppresses floral immunity through promoting GA biosynthesis by the effector SCRE9

Rice false smut caused by Ustilaginoidea virens has become one of the most important rice diseases. U. virens specifically infects rice florets through stamen filaments. However, it is mysterious how U. virens inhibits floral immunity for successful colonization. Here, we show that a small cysteine-rich effector SCRE9 is a key virulence factor in U. virens. Expression of SCRE9 in rice not only suppresses immune responses and false smut resistance, but also significantly increases expression of expansin and gibberellin (GA) biosynthetic genes and GA content in young rice panicles. We further demonstrate that OsSIP1 (SCRE9-interacting protein 1) in rice is targeted by SCRE9 and interacts with the OsMADS63 and OsMADS68 transcription factors that transcriptionally regulate the GA synthesis gene GA3ox1. Interestingly, the chloroplast-localized OsSIP1 recruits OsMADS63/68 into the chloroplast. SCRE9 causes OsSIP1 to be translocated into the nucleus, thereby releasing OsMADS63/68 from the chloroplast into the nucleus to promote GA3ox1 transcription. Therefore, SCRE9 promotes GA biosynthesis and expression of expansins in rice florets, thus loosening cell walls and overcoming the physical barrier during U. virens infection. The finding uncovers an unidentified floret infection strategy that offsets the deficiency in cell wall degrading ability in the unique floret colonizing fungus.

plant biology↗