Research progress on the bulb expansion and starch enrichment in taro (Colocasia esculenta (L). Schott).

Background: Taro is an important potato crop, which can be used as food, vegetable, feed, and industrial raw material. The yield and quality of taro are primarily determined by the expansion degree of taro bulb and the filling condition of starch, whereas the expansion of taro bulb is a complex biological process. However, little information is reviewed on the research progress of bulb expansion and starch enrichment in taro. Methodology: PubMed, Web of Science, and the China National Knowledge Infrastructure databases were searched for relevant articles. After removing duplicate articles and articles with little relevance, 73 articles were selected for review. Results: This article introduces the formation and development of taro bulb for workers engaged in taro research. The content includes the process of amyloplast formation at the cytological level and changes in bulb expansion and starch enrichment at physiological levels, which involve endogenous hormones and key enzyme genes for starch synthesis. The effects of environment and cultivation methods on taro bulb expansion were also reviewed. Conclusions: Future research directions and research focus about the development of taro bulb were proposed. Limited research has been conducted on the physiological mechanism and hormone regulatory pathway of taro growth and development, taro bulb expansion, key gene expression, and starch enrichment. Therefore, the abovementioned research will become the key research direction in the future.

Accumulation and physicochemical properties of starch in relation to eating quality in different parts of taro (Colocasia esculenta) corm.

The accumulation and physicochemical properties of starch affect the eating quality of taro corm. This study aims to investigate the accumulation, morphology, and physicochemical properties of starch from inner and outer tissues in the top, middle, and basal parts of taro corm. Structural and morphological observations showed that the inner tissues of the taro corm accumulated more starch, and the middle tissue had moderate amylose content and the largest granule diameter. Starch from different tissues exhibited A-type orthorhombic structure and similar nuclear magnetic resonance spectrum. The relative crystallinity of starch in the middle tissue was higher than that in the top and basal tissues. Compared with middle and basal tissues, starch from top tissue showed higher peak viscosity, pasting time, swelling power and solubility. Compared with the top and basal tissues, the middle tissue of taro corm exhibited higher index of eating quality including smell, texture, and total evaluation score. The results indicated that starches in various spatial parts of taro corm exhibit differences in accumulation, morphology, structure and physicochemical properties that lead to diverse eating qualities.

The relationship between characteristics of root morphology and grain filling in wheat under drought stress.

Drought is a common yield limiting factor in wheat production and has become a significant threat to global food security. Root system is the organ responsible for water uptake from soil and root growth is closely associated with yield and quality of wheat. However, the relationship between morphological and structural characteristics of root growth and caryopsis enrichment in wheat under drought stress is unclear. In this study, two wheat cultivars (YM13 and YN19) were treated with drought from flowering to caryopsis maturity stage. The changes in morphological structure of roots and characteristics of endosperm enrichment were investigated. Drought stress significantly reduced the root length, plant height, root dry weight and aboveground parts dry weight, whereas the root-shoot ratio of YM13 and YN19 increased by 17.65% and 8.33% under drought stress, respectively. The spike length, spike weight, grains number per spike and 1,000-grains weight of mature wheat also significantly declined under drought stress. Meanwhile, the cross section structure of roots was changed with the enlargement of vascular cylinder and dense distribution of xylem vessels under drought stress. Additionally, drought stress affected the substance enrichment in wheat caryopses, decreasing starch accumulation and increasing protein accumulation of endosperm. Correlation analysis suggested that the root length was closely correlated with the relative areas of amyloplast (0.51) and protein body (0.70), and drought stress increased the correlation coefficient (0.79 and 0.78, respectively). While the root dry weight had a significantly positive correlation with the plant height and aboveground parts dry weight. The results can provide theoretical basis for root architecture optimization, water-saving and high-yield cultivation and quality improvement in wheat.

Analysis of development, accumulation and structural characteristics of starch granule in wheat grain under nitrogen application.

Nitrogen is one of the most important nutrients for wheat growth and has a critical influence on yield and quality. This study aims to examine how medium nitrogen level (240 kg/hm2) affects the starch granule development, starch accumulation, and structural characteristics of wheat starch. The results showed that nitrogen treatment could reduce the biosynthesis of starch and amylose, delay the degradation of starch in pericarp, and promote the proportion of B-type small starch granule in endosperm compared with those in the N0. In addition, the composition and distribution of starch granules were changed, the crystal structure in the inner lamellae and ordered structure in the external region of starch granules were affected, and the swelling power and solubility of starch during wheat development were increased. The effect of nitrogen treatment on the mRNA expression of enzymes related to starch biosynthesis or degradation varied in different developmental stages. During middle and later grain filling stages, AGPase, GBSSI, and GBSSII were lower, and SSS, SBE, and DBE were higher in N240 than in N0. This study indicated that nitrogen application at booting stage significantly affected the structural characteristics of starch, and ultimately determines its quality.

New insights into the mechanism of storage protein biosynthesis in wheat caryopsis under different nitrogen levels.

Effect of different nitrogen levels (0, 150, and 300 kg hm-2) at booting stage on storage protein biosynthesis and processing quality of wheat was investigated using microstructural and ultrastructural observation, RNA sequencing, and quality analysis in this study. The results showed that the storage protein genes encoding ω- and γ-gliadin and low molecular weight glutenin subunit were upregulated at N150, and the genes encoding α- or ß-gliadin and avenin-like protein were upregulated at N300. Two nitrogen levels induced expression of some interesting regulating genes, such as USE1, STX1B_2_3, SEC23, SEC24, SEC61A, HSP A1_8, HSP20, and HSP90B/TRA1. These regulatory genes were enriched in the KEGG pathway protein export, SNARE interactions in vesicular transport, and protein processing in endoplasmic reticulum. The amount, morphology, and accumulation pattern of protein body in four different endosperm regions in developing caryopsis show different response to N150 and N300, of which N300 had greater influence than N150. N150 and N300 both enhanced the contents of protein components, endosperm fullness, grain hardness, and parameters of processing quality, with the latter showing a greater degree of influence. Contrary to the accumulation pattern of protein body, N300 reduced the ratio of the amount of starch granules to the area ratio of protein body to starch granule. Results suggested that the difference of different nitrogen levels affecting storage protein biosynthesis might be through affecting the expression of the encoding and regulating gene of storage protein.

Rice starch accumulation at different endosperm regions and physical properties under nitrogen treatment at panicle initiation stage.

The quality of rice grain is characterized by the component, structure and physicochemical properties of starch accumulated in endosperm cell. Nitrogen uptake strongly affects rice growth and starch development. In this study, Nangeng 9108 was used to investigated the accumulation of starch in different positions of the endosperm and physical properties of starch under nitrogen treatment of panicle initiation (PI) stage. Compared with the control group (CG), nitrogen treatment group (NTG) featured a higher number of grains per panicle and 1000-grain weight. Nitrogen treatment significantly increased starch accumulation among different regions during endosperm development, which was expressed as central endosperm cells > sub-aleurone cells of abdominal endosperm > sub-aleurone cells of dorsal endosperm. The amyloplast increased by constricting and budding-type division, generated a bead-like structure and derived some vesicles. The particle size of the starch granules obtained from the NTG was smaller and the apparent amylose content was lower than those of the CG, resulting in higher relative crystallinity. Nitrogen treatment promoted double helical components and provided a higher degree of order at short-rang scale for the starch granules. This study indicated that nitrogen significantly affected the accumulation and physicochemical properties of starch in the endosperm.

Exogenous application of NaBiF4 nanoparticle affects wheat root development.

BACKGROUND: Nanoparticle causes soil pollution, which affected plant development and then resulted in biomass decreased, especially in crops. However, little is known how sodium nanoparticles affect wheat root development at plant physiological level. RESULTS: We used NaBiF4 (size of 50-100 nm) to analyze the effect in wheat development at plant physiological level. Under exogenous application of 50 µM NaBiF4 for treatment, wheat root elongation was inhibited, but fresh weight and dry weight were increased. We also found that NaBiF4 induced that the plant had lower content of sodium than negative control. Used no-sodium nanoparticle of BiF3 for another negative control, it was also supported that NaBiF4 entered into cell to replace of sodium and exported sodium out of plant. These results implied NaBiF4 might induce sodium export to maintain the balance between sodium and potassium elements. Additionally, metabolism analysis demonstrated that SOD activity was increased, but CAT and POD activity reduced under exogenous treatment of NaBiF4 nanoparticles. CONCLUSIONS: Sodium nanoparticles (NaBiF4) inhibited plant development by nanoparticle accumulation and sodium homeostasis broken, and then involved reactive oxygen species (ROS) signaling system response. These results provided more sights of sodium nanoparticle effect in plant development.

Novel insights into the effect of drought stress on the development of root and caryopsis in barley.

Drought is a common natural disaster in barley production, which restricts the growth and development of barley roots and caryopses seriously, thereby decreasing yield and debasing grain quality. However, mechanisms for how drought stress affects barley caryopses and roots development under drought stress are unclear. In this paper, Suluomai1 was treated with drought from flowering to caryopses mature stage. The morphological and structural changes in roots growth and caryopses development of barley were investigated. Drought stress increased root/shoot ratio and eventually led to the 20.16% reduction of ear weight and 7.75% reduction of 1,000-grain weight by affecting the biomass accumulation of roots and caryopses. The barley roots under drought had more lateral roots while the vessel number and volume of roots decreased. Meanwhile, drought stress accelerated the maturation of caryopses, resulting in a decrease in the accumulation of starch but a significant increase of protein accumulation in barley endosperm. There was a significantly positive correlation (0.76) between the area of root vessel and the relative area of protein in endosperm cells under normal condition and drought increased the correlation coefficient (0.81). Transcriptome analysis indicated that drought induced differential expressions of genes in caryopses were mainly involved in encoding storage proteins and protein synthesis pathways. In general, drought caused changes in the morphology and structure of barley roots, and the roots conveyed stress signals to caryopses, inducing differential expression of genes related to protein biosynthesis, ultimately leading to the increase in the accumulation of endosperm protein. The results not only deepen the study on drought mechanism of barley, but also provide theoretical basis for molecular breeding, high-yield cultivation and quality improvement in barley.

Effects of high temperature during two growth stages on caryopsis development and physicochemical properties of starch in rice.

Global warming may affect the development of rice at different growth stages, thereby decreasing rice yield and deteriorating grain quality. The difference in rice responses to high temperature during primordial differentiation (PD) and pollen filling (PF) stages has been rarely studied. In this paper, two temperature treatments (40 °C and 30 °C) at the two stages (PD and PF) were imposed to four rice groups under the controlled temperature chambers. Compared with rice under normal temperature, high temperature-stressed rice showed accelerated growth rate, smaller caryopsis and decreased yield. Moreover, high temperature affected the starch physicochemical properties, resulting in lower apparent amylose content and higher order degree, gelatinization temperatures, and thereby increased peak, trough and final viscosities in starch. High temperature during PD stage inhibited cell development and starch deposition, thus leading to small starch granule and low retrogradation. However, temperature-stressed rice during PF stage showed increased starch accumulation and larger granule size. Therefore, effects of high temperature during the two stages on caryopsis development and starch properties were partly similar but also notably different. These results enriched and deepened the study of high temperature-stressed rice and served as an important reference for the processing and utilization of rice starch in food industry.

Structural development and physicochemical properties of starch in caryopsis of super rice with different types of panicle.

BACKGROUND: Starch is the main storage substance in rice caryopsis and its properties will determine the quality of rice. Super rice has been extensively studied due to its high-yield characteristics, but the knowledge of amyloplast development and starch quality in caryopsis of super rice especially with large panicle is limited. RESULTS: To address this, large panicle typed and normal panicle typed super rice cultivar Yongyou2640 (YY2640) and Nangeng9108 (NG9108) were investigated in this study. The development of amyloplast in YY2640 caryopsis was better than NG9108, showing faster degradation rate of pericarp amyloplast and better filling degree of endosperm amyloplast. Meanwhile, the starch granule of YY2640 presented as polyhedral shape with smooth surface and the granule size was slightly larger than NG9108. The starch of YY2640 exhibited the lower amylose content, ratio of amylose to amylopectin and the higher level of amylopectin short and long branch-chains compared with NG9108, but there was no significant difference in amylopectin branching degree between them. Two rice starches both showed the characteristics of A-type crystal, and the relative crystallinity and external ordered degree of YY2640 starch were higher than those of NG9108. Furthermore, YY2640 starch showed better pasting properties with lower pasting temperature, shorter pasting time, higher peak viscosity, trough viscosity, breakdown value and lower setback value because of lower apparent amylose content. CONCLUSIONS: Overall, the development and filling of amyloplast in YY2640 caryopsis were better than those of NG9108, thus leading to better starch quality of YY2640.

Novel Insights into miRNA Regulation of Storage Protein Biosynthesis during Wheat Caryopsis Development under Drought Stress.

Drought stress is a significant abiotic stress factor that affects wheat yield and quality. MicroRNA (miRNA) plays an important role in regulating caryopsis development in response to drought stress. However, little is known about the expression characteristics of miRNAs and how they regulate protein accumulation in wheat caryopsis under drought stress. To address this, two small RNA libraries of wheat caryopsis under control and drought stress conditions were constructed and sequenced. A total of 125 miRNAs were identified in the two samples, of which 110 were known and 15 were novel. A total of 1,981 miRNA target genes were predicted and functional annotations were obtained from various databases for 1,641 of them. Four miRNAs were identified as differential expression under drought stress, and the expression patterns of three of them were consistent with results obtained by reverse transcription polymerase chain reaction (RT-PCR) and reverse transcription quantitative polymerase chain reaction (RT-qPCR). Moreover, three miRNA-target pairs showed negative regulation tendency, as revealed by RT-qPCR. Functional enrichment and pathway analysis revealed that four pathways might be involved in storage protein biosynthesis. Furthermore, drought stress significantly increased the accumulation of protein bodies and protein content in wheat endosperm. In summary, our findings suggest that drought stress may enhance storage protein by regulating the expression of miRNAs and their target genes.

Novel insights into the effect of nitrogen on storage protein biosynthesis and protein body development in wheat caryopsis.

Molecular and cytological mechanisms concerning the effects of nitrogen on wheat (Triticum aestivum L.) storage protein biosynthesis and protein body development remain largely elusive. We used transcriptome sequencing, proteomics techniques, and light microscopy to investigate these issues. In total, 2585 differentially expressed genes (DEGs) and 57 differentially expressed proteins (DEPs) were found 7 days after anthesis (DAA), and 2456 DEGs and 64 DEPs were detected 18 DAA after nitrogen treatment. Gene ontology terms related to protein biosynthesis processes enriched these numbers by 678 and 582 DEGs at 7 and 18 DAA, respectively. Further, 25 Kyoto Encyclopedia of Genes and Genomes pathways were involved in protein biosynthesis at both 7 and 18 DAA. DEPs related to storage protein biosynthesis contained gliadin and glutenin subunits, most of which were up-regulated after nitrogen treatment. Quantitative real-time PCR analysis indicated that some gliadin and glutenin subunit encoding genes were differentially expressed at 18 DAA. Structural observation revealed that wheat endosperm accumulated more and larger protein bodies after nitrogen treatment. Collectively, our findings suggest that nitrogen treatment enhances storage protein content, endosperm protein body quantity, and partial processing quality by altering the expression levels of certain genes involved in protein biosynthesis pathways and storage protein expression at the proteomics level.

Regional analysis of potential polychlorinated biphenyl degrading bacterial strains from China

ABSTRACT Polychlorinated biphenyls (PCBs), the chlorinated derivatives of biphenyl, are one of the most prevalent, highly toxic and persistent groups of contaminants in the environment. The objective of this study was to investigate the biodegradation of PCBs in northeastern (Heilongjiang Province), northern (Shanxi Province) and eastern China (Shanghai municipality). From these areas, nine soil samples were screened for PCB-degrading bacteria using a functional complementarity method. The genomic 16S rDNA locus was amplified and the products were sequenced to identify the bacterial genera. Seven Pseudomonas strains were selected to compare the capacity of bacteria from different regions to degrade biphenyl by HPLC. Compared to the biphenyl content in controls of 100%, the biphenyl content went down to 3.7% for strain P9-324, 36.3% for P2-11, and 20.0% for the other five strains. These results indicate that a longer processing time led to more degradation of biphenyl. PCB-degrading bacterial strains are distributed differently in different regions of China.

Regional analysis of potential polychlorinated biphenyl degrading bacterial strains from China.

Polychlorinated biphenyls (PCBs), the chlorinated derivatives of biphenyl, are one of the most prevalent, highly toxic and persistent groups of contaminants in the environment. The objective of this study was to investigate the biodegradation of PCBs in northeastern (Heilongjiang Province), northern (Shanxi Province) and eastern China (Shanghai municipality). From these areas, nine soil samples were screened for PCB-degrading bacteria using a functional complementarity method. The genomic 16S rDNA locus was amplified and the products were sequenced to identify the bacterial genera. Seven Pseudomonas strains were selected to compare the capacity of bacteria from different regions to degrade biphenyl by HPLC. Compared to the biphenyl content in controls of 100%, the biphenyl content went down to 3.7% for strain P9-324, 36.3% for P2-11, and 20.0% for the other five strains. These results indicate that a longer processing time led to more degradation of biphenyl. PCB-degrading bacterial strains are distributed differently in different regions of China.

Comparison of Morphology and Physicochemical Properties of Starch Among 3 Arrowhead Varieties.

Arrowhead (Sagittaria trifolia var. sinensis) is a source of starch worldwide, but arrowhead starch has been rarely studied. In this work, starch was separated from arrowhead corm. The morphology and physicochemical properties of starch were then investigated and compared among 3 different arrowhead varieties (Purple-corm, Hongta, and Japanese). Results showed that starches from the 3 varieties similarly featured an oval shape containing a visible polarization cross, a CA -type crystalline structure, and an ordered structure in the external granule region. However, starch content, granule size, crystal characteristics, and pasting properties differed among the 3 varieties. Japanese arrowhead exhibited the highest starch content and degree of ordered structure in the external granule region, as well as onset, peak, and final gelatinization temperature. Purple-corm arrowhead starch demonstrated the highest amylose content and relative degree of crystallinity, smallest granule size, and lowest swelling power and solubility. Purple-corm arrowhead starch also showed the highest gelatinization enthalpy, as well as peak, trough, final, and setback viscosities. This starch further presented the lowest breakdown viscosity and degree of hydrolysis by HCl and porcine pancreatic α-amylase. These findings can provide useful references for arrowhead variety selection in food and nonfood industries.

Effect of drought stress on the development of endosperm starch granules and the composition and physicochemical properties of starches from soft and hard wheat.

BACKGROUND: The objectives of this study were to: (1) observe the effects of drought stress (DS) on the structural development of endosperm starch granules; (2) investigate the effects of DS on composition and physicochemical properties of starches; and (3) compare the different responses to DS between soft and hard wheat. RESULTS: DS resulted in large A-type starch granules at 12 d after anthesis (DAA) and a high percentage of B-type starch granules at 18 DAA in endosperm cells of the two wheat cultivars. DS decreased the 1000-grain weight, total starch and amylose contents, and amylose-to-amylopectin ratio of both starches. DS also decreased the percentage of B-type starch granules in NM13 and increased the number of hollows on the surface of A-type starch granules in XM33. DS further increased the swelling power and affected pasting properties of both starches. DS also significantly enhanced the hydrolysis degrees of starches by pancreatic α-amylase, Aspergillus niger amyloglucosidase, and HCl in NM13. DS altered the contents of rapidly digestible, slowly digestible, and resistant starches in native, gelatinised, and retrograded starches. CONCLUSION: Overall, DS can affect the development of endosperm starch granules and the physicochemical properties of starches, thus affecting the qualities of the final wheat products. © 2015 Society of Chemical Industry.

Physicochemical Properties of Starch Isolated from Bracken (Pteridium aquilinim) Rhizome.

Bracken (Pteridium aquilinum) is an important wild plant starch resource worldwide. In this work, starch was separated from bracken rhizome, and the physicochemical properties of this starch were systematically investigated and compared with 2 other common starches, that is, starches from waxy maize and potato. There were significant differences in shape, birefringence patterns, size distribution, and amylose content between bracken and the 2 other starches. X-ray diffraction analysis revealed that bracken starch exhibited a typical C-type crystalline structure. Bracken starch presented, respectively, lower and higher relative degree of crystallinity than waxy maize and potato starches. Ordered structures in particle surface differed among these 3 starches. The swelling power tendency of bracken starch in different temperature intervals was very similar to that of potato starch. The viscosity parameters during gelatinization were the lowest in waxy maize, followed by bracken and potato starches. The contents of 3 nutritional components, that is, rapidly digestible, slowly digestible, and resistant starches in native, gelatinized, and retrograded starch from bracken rhizome presented more similarities with potato starch than waxy maize starch. These finding indicated that physicochemical properties of bracken starch showed more similarities with potato starch than waxy maize starch.

Systematic Analysis of Pericarp Starch Accumulation and Degradation during Wheat Caryopsis Development.

Although wheat (Triticum aestivum L.) pericarp starch granule (PSG) has been well-studied, our knowledge of its features and mechanism of accumulation and degradation during pericarp growth is poor. In the present study, developing wheat caryopses were collected and starch granules were extracted from their pericarp to investigate the morphological and structural characteristics of PSGs using microscopy, X-ray diffraction and Fourier transform infrared spectroscopy techniques. Relative gene expression levels of ADP-glucose pyrophosphorylase (APGase), granule-bound starch synthase II (GBSS II), and α-amylase (AMY) were quantified by quantitative real-time polymerase chain reaction. PSGs presented as single or multiple starch granules and were synthesized both in the amyloplast and chloroplast in the pericarp. PSG degradation occurred in the mesocarp, beginning at 6 days after anthesis. Amylose contents in PSGs were lower and relative degrees of crystallinity were higher at later stages of development than at earlier stages. Short-range ordered structures in the external regions of PSGs showed no differences in the developing pericarp. When hydrolyzed by α-amylase, PSGs at various developmental stages showed high degrees of enzymolysis. Expression levels of AGPase, GBSS II, and AMY were closely related to starch synthesis and degradation. These results help elucidate the mechanisms of accumulation and degradation as well as the functions of PSG during wheat caryopsis development.

Morphology and Physicochemical Properties of 3 Lilium Bulb Starches.

Lilium (Liliaceae) is an important wild plant and is used as food and traditional medicine worldwide. One Lilium cultivar (Lilium lancifolium) and 2 wild types (Lilium leucanthum and Lilium rosthornii) that are commonly distributed in Western China were investigated to completely utilize Lilium resources. The morphology of the flowers, bulbs, and scales and soluble sugar, total starch and amylose contents was remarkably different among the 3 Lilium species. Starches from the 3 Lilium species presented different granule size and shape. The starch of L. lancifolium exhibited higher swelling power and solubility than that of L. leucanthum and L. rosthornii. The starches from the 3 Lilium bulbs presented similar X-ray diffraction patterns and Fourier transform infrared spectroscopy. Among the 3 Lilium species, L. lancifolium showed the lowest crystallinity and the largest proportion of ordered structures in granule external region. Gelatinization temperatures and retrogradation percentage were significantly lower, but gelatinization enthalpy was significantly higher in L. lancifolium than those in L. leucanthum and L. rosthornii. Pasting properties of starch were different among the 3 Lilium species. Starch from L. lancifolium showed the highest degree of amylopectin branching, followed by L. leucanthum and L. rosthornii. Starches from L. leucanthum and L. rosthornii showed higher resistance to porcine pancreatic α-amylase hydrolysis compared to that of L. lancifolium. These results indicated that 3 Lilium bulbs exhibited remarkable differences in morphological, crystal, thermal, pasting, and hydrolysis properties of starches.

De novo assembly, transcriptome characterization, lignin accumulation, and anatomic characteristics: novel insights into lignin biosynthesis during celery leaf development.

Celery of the family Apiaceae is a biennial herb that is cultivated and consumed worldwide. Lignin is essential for cell wall structural integrity, stem strength, water transport, mechanical support, and plant pathogen defense. This study discussed the mechanism of lignin formation at different stages of celery development. The transcriptome profile, lignin distribution, anatomical characteristics, and expression profile of leaves at three stages were analyzed. Regulating lignin synthesis in celery growth development has a significant economic value. Celery leaves at three stages were collected, and Illumina paired-end sequencing technology was used to analyze large-scale transcriptome sequences. From Stage 1 to 3, the collenchyma and vascular bundles in the petioles and leaf blades thickened and expanded, whereas the phloem and the xylem extensively developed. Spongy and palisade mesophyll tissues further developed and were tightly arranged. Lignin accumulation increased in the petioles and the mesophyll (palisade and spongy), and the xylem showed strong lignification. Lignin accumulation in different tissues and at different stages of celery development coincides with the anatomic characteristics and transcript levels of genes involved in lignin biosynthesis. Identifying the genes that encode lignin biosynthesis-related enzymes accompanied by lignin distribution may help elucidate the regulatory mechanisms of lignin biosynthesis in celery.