The DOI matches its Crossref record; 60 of the 62 references it cites were found in a registry or an index.
Lignin and Biomass: A Negative Correlation for Wood Formation and Lignin Content in Trees
Open accessAbstract
Studies in populations of forest tree hybrids have shown a negative correlation of biomass growth (usually measured as wood volume) and lignin content (Kirst et al., 2004; Novaes et al., 2009). The control of growth and lignin appears to be highly regulated, implying that selection for improved growth rate could effect a reduction in lignin content. Trees with increased biomass and reduced lignin would provide a yield advantage for pulp and paper production, as well as for production of biofuels. The purpose of this article is to briefly review the evidence for the correlation between lignin content and biomass growth and to discuss this correlation in a metabolic context. We review evidence that the regulation of the balance between lignin biosynthesis and biomass growth is mediated, at least in part, at the level of transcription, and expand previous results to show specific regulation within gene families of monolignol biosynthesis. Finally, we suggest a model of physiological control for the regulation of the relationship of wood formation and lignin content in trees. Wood, used by human societies for millennia, remains one of the world’s most abundant raw materials for industrial products and renewable energy. Wood is the secondary xylem of vascular plants, a tissue formed from the terminal differentiation of the inner side of the cambial meristem for vertical and horizontal transport of water, nutrients, and extractives. The secondary cell wall structure and composition of wood are the primary determinants of its physical and chemical properties, and of its energy content. Wood is typically composed of about 25% lignin, and 70% cellulosic carbohydrates, with roughly 45% cellulose and 25% hemicelluloses (Sjostrom, 1993). Lignin is a complex phenolic polymer that provides an embedding material for the cellulosic polymers of the secondary cell walls. It is also the major polymer in the middle lamellae between adjacent cell walls (Plomion et al., 2001; Boerjan et al., 2003). Lignin provides the hydrophobic surface that allows plants to transport water to heights greater than 100 m (Carder, 1995; Koch et al., 2004) and contributes to the mechanical strength that can support trees weighing more than 2,000 metric tons (Fry and White, 1938). Lignin’s physical and chemical properties also serve as a barrier against the invasion of pests and pathogens (Vance et al., 1980; Bhuiyan et al., 2009). For the forest products industries, lignin is the major barrier to efficient extraction of cellulose fibers for pulp and paper production. For the bioenergy industries, lignin is a barrier to saccharification for production of liquid biofuel (Li et al., 2003; Chen and Dixon, 2007). While lignin is fundamental to growth and adaptation of herbaceous and woody plants (Sarkanen and Ludwig, 1971; Boerjan et al., 2003; Ralph et al., 2007; Heitner et al., 2010; Vanholme et al., 2010), it is not clear how much lignin is needed or how much its composition may vary. Within a plant, lignin content can vary greatly in different tissues; for example, lignin is very low in young shoots and high in wood. In different tree species the lignin content of wood can vary from 15% to 40% (Sarkanen and Ludwig, 1971). Within a species, average lignin content of wood is much less variable, often ranging only a few percent (Einspahr et al., 1964; van Buijtenen et al., 1968). Lignin content is also increased or decreased in wood formed under gravitropic stimulation or mechanical stress (known as reaction wood; Timell, 1969, 1986; Andersson-Gunneras et al., 2006). In softwoods (gymnosperms), compression wood may be up to 40% lignin, and in hardwoods (angiosperms), tension wood fibers have a specialized gelatinous cell layer that is almost devoid of lignin (Timell, 1969). Lignin composition is also highly variable. There are two main types of subunits in lignin. One derived from coniferyl alcohol monomers forms guaiacyl (G) units, resulting in a lignin typical of softwoods. A second monomer sinapyl alcohol forms syringyl (S) units after polymerization. A mixed type of lignin with both G and S units is typical of hardwoods, and is characterized by the ratio (S/G ratio) of subunits. The S/G ratio varies greatly among hardwoods (Sarkanen and Ludwig, 1971). In herbaceous and field crops, significant reductions in lignin have been achieved by traditional breeding, spontaneous mutation, and by transgenesis. In general, reduced lignin is associated with depressed yields (Pedersen et al., 2005). However, in many cases, lignin can be reduced without reducing yield or fitness. Genetic background and diverse environments have a significant role. Because of its roles in water transport, mechanical support, and biodefense, lignin is intimately associated with the evolution of vascular plants (Robinson, 1990). Early large plants may have had higher lignin contents than we find today. Ferns and lycopods, the predominant plants of the middle to late Paleozoic (350–
Overview
The claims and statistics of this paper haven’t been checked yet.
Run the check — free, no account needed. It reads the open-access text and checks the sources it cites.
Integrity signals
Registry
Cites
Statistics
Verification
Claims the paper makes
The claims and statistics of this paper haven’t been checked yet.
Run the check — free, no account needed. It reads the open-access text and checks the sources it cites.
Claims other papers make about this one
Each is checked against this paper’s own text: does it say what the citing paper says it does?
The source backs part of the claim, not all of it — or backs it for a different population, condition or comparison.
In the citing paper
“It seems likely that at first methoxy groups are cleaved by the high temperature and acidity of the catalyst. As most of the monomers of lignins consist of syringyl (S) and guaiacyl units (G) and to a lesser extent of p-hydroxphenyl units (H), this results in a mixture of mainly catechols/pyrogallols.39”
From Synthesis of Bio-aromatics from Black Liquors Using Catalytic Pyrolysis. (2018) · cites this paper as [39]
Experimental — accuracy not yet measured
Self-check
Percentages, sums and reported differences re-computed from the paper’s own numbers.
The self-check needs the paper’s full text.
Citations
Cites
Nutritional ecology of the ruminant
Soest, P. J. V. (1995) · Choice Reviews Online · doi:10.5860/choice.32-4505 · Cited by 7,714
Boerjan, W., Ralph, J., & Baucher, M. (2003) · Annual Review of Plant Biology · doi:10.1146/annurev.arplant.54.031902.134938 · Cited by 4,504
Wood: Chemistry, ultrastructure, reactions
(1983) · doi:10.1515/9783110839654 · Cited by 2,846
Wood-chemistry, ultrastructure, reactions
Nimz, H. (1984) · European Journal of Wood and Wood Products · doi:10.1007/bf02608943 · Cited by 2,774
Lignin Biosynthesis and Structure
Vanholme, R., Demedts, B., Morreel, K., Ralph, J., & Boerjan, W. (2010) · PLANT PHYSIOLOGY · doi:10.1104/pp.110.155119 · Cited by 2,686
Lignins. Occurrence, Formation, Structure and Reactions
Sarkanen, K. V., & Ludwig, C. H. (1971) · Medical Entomology and Zoology · Cited by 2,008
The phenylpropanoid pathway and plant defence—a genomics perspective
Dixon, R. A., Achnine, L., Kota, P., Liu, C., Reddy, M. S. S., & Wang, L. (2002) · Molecular Plant Pathology · doi:10.1046/j.1364-3703.2002.00131.x · Cited by 1,494
Sucrose metabolism: regulatory mechanisms and pivotal roles in sugar sensing and plant development
Koch, K. E. (2004) · Current Opinion in Plant Biology · doi:10.1016/j.pbi.2004.03.014 · Cited by 1,487
Lignin modification improves fermentable sugar yields for biofuel production
Chen, F., & Dixon, R. A. (2007) · Nature Biotechnology · doi:10.1038/nbt1316 · Cited by 1,347
Koch, G. W., Sillett, S. C., Jennings, G. M., & Davis, S. D. (2004) · Nature · doi:10.1038/nature02417 · Cited by 1,114
Scheible, W., Morcuende, R., Czechowski, T., Fritz, C., Osuna, D., Palacios‐Rojas, N., Schindelasch, D., Thimm, O., Udvardi, M. K., & Stitt, M. (2004) · PLANT PHYSIOLOGY · doi:10.1104/pp.104.047019 · Cited by 1,047
Lignification as a Mechanism of Disease Resistance
Vance, C. P., Kirk, T. K., & Sherwood, R. T. (1980) · Annual Review of Phytopathology · doi:10.1146/annurev.py.18.090180.001355 · Cited by 975
Zhou, J., Lee, C., Zhong, R., & Ye, Z. (2009) · The Plant Cell · doi:10.1105/tpc.108.063321 · Cited by 869
Compression Wood in Gymnosperms
Timell, T. E. (1986) · Springer eBooks · doi:10.1007/978-3-642-61616-7 · Cited by 851
Genome-Wide Characterization of the Lignification Toolbox in Arabidopsis
Raes, J., Rohde, A., Christensen, J. H., de Peer, Y. V., & Boerjan, W. (2003) · PLANT PHYSIOLOGY · doi:10.1104/pp.103.026484 · Cited by 835
Repression of lignin biosynthesis promotes cellulose accumulation and growth in transgenic trees
Hu, W., Harding, S. A., Lung, J., Popko, J. L., Ralph, J., Stokke, D. D., Tsai, C., & Chiang, V. L. (1999) · Nature Biotechnology · doi:10.1038/11758 · Cited by 794
Self-citationPlomion, C., Provost, G. L., & Stokes, A. (2001) · PLANT PHYSIOLOGY · doi:10.1104/pp.010816 · Cited by 707
Plomion, C., Leprovost, G., & Stokes, A. (2001) · PubMed · Cited by 663
Sederoff, H. W., & Huber, S. C. (2000) · Critical Reviews in Biochemistry and Molecular Biology · doi:10.1080/10409230008984165 · Cited by 627
Tamagnone, L., Mérida, Á., PARR, A. J., Mackay, S., Culiáñez‐Macià, F. A., Roberts, K., & Martin, C. R. (1998) · The Plant Cell · doi:10.1105/tpc.10.2.135 · Cited by 547
Riou, C., Menges, M., Healy, J. M. S., & Murray, J. A. H. (2000) · Molecular and Cellular Biology · doi:10.1128/mcb.20.13.4513-4521.2000 · Cited by 440
Combinatorial modification of multiple lignin traits in trees through multigene cotransformation
Li, L., Zhou, Y., Cheng, X., Sun, J., Marita, J. M., Ralph, J., & Chiang, V. L. (2003) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.0831166100 · Cited by 425
Self-citationColeman, H. D., Yan, J., & Mansfield, S. D. (2009) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.0900188106 · Cited by 416
Fritz, C., Palacios‐Rojas, N., Feil, R., & Stitt, M. (2006) · The Plant Journal · doi:10.1111/j.1365-313x.2006.02715.x · Cited by 403
Andersson‐Gunnerås, S., Mellerowicz, E. J., Love, J., Segerman, B., Ohmiya, Y., Coutinho, P. M., Nilsson, P., Henrissat, B., Möritz, T., & Sundberg, B. (2005) · The Plant Journal · doi:10.1111/j.1365-313x.2005.02584.x · Cited by 397
Goicoechea, M., Lacombe, E., Legay, S., Mihaljević, S., Rech, P., Jauneau, A., Lapierre, C., Pollet, B., Verhaegen, D., Chaubet, N., et al. (2005) · The Plant Journal · doi:10.1111/j.1365-313x.2005.02480.x · Cited by 369
Kammerer, B., Fischer, K., Hilpert, B., Schubert, S., Gutensohn, M., Weber, A. P. M., & Flügge, U. (1998) · The Plant Cell · doi:10.1105/tpc.10.1.105 · Cited by 335
Role of lignification in plant defense
Bhuiyan, N. H., Selvaraj, G., Wei, Y., & King, J. (2009) · Plant Signaling & Behavior · doi:10.4161/psb.4.2.7688 · Cited by 326
Shi, R., Sun, Y., Li, Q., Heber, S., Sederoff, R. R., & Chiang, V. L. (2009) · Plant and Cell Physiology · doi:10.1093/pcp/pcp175 · Cited by 322
Self-citationIndole-3-Acetic Acid Controls Cambial Growth in Scots Pine by Positional Signaling1
Uggla, C., Mellerowicz, E. J., & Sundberg, B. (1998) · PLANT PHYSIOLOGY · doi:10.1104/pp.117.1.113 · Cited by 314
Functional Characterization of Poplar Wood-Associated NAC Domain Transcription Factors
Zhong, R., Lee, C., & Ye, Z. (2009) · PLANT PHYSIOLOGY · doi:10.1104/pp.109.148270 · Cited by 292
Biochemistry and Molecular Biology of Wood
Higuchi, T. (1997) · Springer series in wood science · doi:10.1007/978-3-642-60469-0 · Cited by 268
Impact of Reduced Lignin on Plant Fitness
Pedersen, J. F., Vogel, K. P., & Funnell, D. L. (2005) · Crop Science · doi:10.2135/cropsci2004.0155 · Cited by 240
EFFECT OF LIGNIN CONTENT AND EXTRACTIVES ON THE HIGHER HEATING VALUE OF WOOD
White, R. H. (1987) · Wood and Fiber Science (Society of Wood Science and Technology) · Cited by 238
Mele, G., Ori, N., Sato, Y., & Hake, S. (2003) · Genes & Development · doi:10.1101/gad.1120003 · Cited by 226
Kirst, M., Myburg, A. A., León, J. P. G. D., Kirst, M. E., Scott, J. T., & Sederoff, R. R. (2004) · PLANT PHYSIOLOGY · doi:10.1104/pp.103.037960 · Cited by 220
Self-citationTrethewey, R. N., Geigenberger, P., Riedel, K., Hajirezaei, M., Sonnewald, U., Stitt, M., Riesmeier, J., & Willmitzer, L. (1998) · The Plant Journal · doi:10.1046/j.1365-313x.1998.00190.x · Cited by 216
Bomal, C., Bedon, F., Caron, S., Mansfield, S. D., Levasseur, C., Cooke, J. E. K., Blais, S., Tremblay, L., Morency, M., Pavy, N., et al. (2008) · Journal of Experimental Botany · doi:10.1093/jxb/ern234 · Cited by 195
Dauwe, R., Morreel, K., Goeminne, G., Gielen, B., Rohde, A., Beeumen, J. V., Ralph, J., Boudet, A. M., Kopka, J., Rochange, S. F., et al. (2007) · The Plant Journal · doi:10.1111/j.1365-313x.2007.03233.x · Cited by 190
Beemster, G. T. S., Vusser, K. D., Tavernier, E. D., Bock, K. D., & Inzé, D. (2002) · PLANT PHYSIOLOGY · doi:10.1104/pp.002923 · Cited by 182
Efficiency of Lignin Biosynthesis: a Quantitative Analysis
Amthor, J. S. (2003) · Annals of Botany · doi:10.1093/aob/mcg073 · Cited by 179
Functional analysis of tobacco LIM protein Ntlim1 involved in lignin biosynthesis
Kawaoka, A., Kaothien-Nakayama, P., Yoshida, K., Endo, S., Yamada, K., & Ebinuma, H. (2000) · The Plant Journal · doi:10.1046/j.1365-313x.2000.00737.x · Cited by 178
McKibbin, R. S., Muttucumaru, N., Paul, M. J., Powers, S. J., Burrell, M. M., Coates, S. A., Purcell, P. C., Tiessen, A., Geigenberger, P., & Halford, N. G. (2006) · Plant Biotechnology Journal · doi:10.1111/j.1467-7652.2006.00190.x · Cited by 169
Novaes, E., Osorio, L. F., Drost, D. R., Miles, B. L., Novaes, C. R. D. B., Benedict, C., Dervinis, C., Yu, Q., Sykes, R. W., Davis, M. F., et al. (2009) · New Phytologist · doi:10.1111/j.1469-8137.2009.02785.x · Cited by 155
Self-citationTranscriptional Control of Monolignol Biosynthesis in Pinus taeda
Anterola, A. M., Jeon, J.-H., Davin, L. B., & Lewis, N. (2002) · Journal of Biological Chemistry · doi:10.1074/jbc.m112051200 · Cited by 135
Tenhaken, R., & Thulke, O. (1996) · PLANT PHYSIOLOGY · doi:10.1104/pp.112.3.1127 · Cited by 128
The failure of nitrogen and lignin control of decomposition in a North American desert
Schaefer, D. A., Steinberger, Y., & Whitford, W. G. (1985) · Oecologia · doi:10.1007/bf00378913 · Cited by 125
Carbon metabolite sensing and signalling
Halford, N. G., & Paul, M. J. (2003) · Plant Biotechnology Journal · doi:10.1046/j.1467-7652.2003.00046.x · Cited by 124
Aloni, R., Tollier, M. T., & Monties, B. B. (1990) · PLANT PHYSIOLOGY · doi:10.1104/pp.94.4.1743 · Cited by 108
98/03070 Evaluating physical, chemical, and energetic properties of perennial grasses as biofuels
(1998) · Fuel and Energy Abstracts · doi:10.1016/s0140-6701(98)96595-x · Cited by 102
Merchant, A., Peuke, A. D., Keitel, C., Macfarlane, C., Warren, C. R., & Adams, M. A. (2010) · Journal of Experimental Botany · doi:10.1093/jxb/erq045 · Cited by 80
Konishi, T., Ohmiya, Y., & Hayashi, T. (2004) · PLANT PHYSIOLOGY · doi:10.1104/pp.103.033167 · Cited by 69
Drost, D. R., Benedict, C. I., Berg, A. S., Novaes, E., Novaes, C. R. D. B., Yu, Q., Dervinis, C., Maia, J. M., Yap, J. S. F., Miles, B. L., et al. (2010) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.0914709107 · Cited by 63
Self-citationCouturier, J., de Faÿ, E., Fitz, M., Wipf, D., Blaudez, D., & Chalot, M. (2010) · Journal of Experimental Botany · doi:10.1093/jxb/erq036 · Cited by 60
Average effect of a mutation in lignin biosynthesis in loblolly pine
Wu, R., Remington, D. L., MacKay, J. J., McKeand, S. E., & O'Malley, D. M. (1999) · Theoretical and Applied Genetics · doi:10.1007/s001220051287 · Cited by 46
Drost, D. R., Novaes, E., Novaes, C. R. D. B., Benedict, C. I., Brown, R. S., Yin, T., Tuskan, G. A., & Kirst, M. (2009) · The Plant Journal · doi:10.1111/j.1365-313x.2009.03828.x · Cited by 46
Self-citationHarding, S. A., Jarvie, M., Lindroth, R. L., & Tsai, C. (2009) · Journal of Experimental Botany · doi:10.1093/jxb/erp180 · Cited by 42
Yu, Q., Li, B., Nelson, C. D., McKeand, S. E., Batista, V. B., & Mullin, T. J. (2006) · Tree Genetics & Genomes · doi:10.1007/s11295-005-0032-y · Cited by 31
Okamoto, H. (1965) · Journal of the Society of Materials Science Japan · doi:10.2472/jsms.14.685 · Cited by 22
Best, J. A., Barazangi, M., Al-Saad, D., Sawaf, T., & Gebran, A. (1990) · Geology · doi:10.1130/0091-7613(1990)018
Flagged first, then by citations
2 of the 62 references couldn’t be matched to a known paper.
Research works printed but not linked: 3
- Einspahr, 1964, Slash pine wood and fiber property heritability study, Silvae Genet, 13, 103
- Timell, 1969, The chemical composition of tension wood, Svensk Papperstidning, 72, 173
- van Buijtenen, 1968, Micropulping loblolly pine grafts selected for extreme wood specific gravity, Silvae Genet, 17, 15
Cited by
Sorted by citations
An overview of the organic and inorganic phase composition of biomass
Vassilev, S. V., Baxter, D., Andersen, L. K., Vassileva, C. G., & Morgan, T. J. (2011) · Fuel · doi:10.1016/j.fuel.2011.09.030 · Cited by 1,085
Lignin content in natural Populus variants affects sugar release
Studer, M. H., DeMartini, J. D., Davis, M. F., Sykes, R. W., Davison, B. H., Keller, M., Tuskan, G. A., & Wyman, C. E. (2011) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.1009252108 · Cited by 588
In Posidonia oceanica cadmium induces changes in DNA methylation and chromatin patterning
Greco, M., Chiappetta, A. A. C., Bruno, L., & Bitonti, M. B. (2011) · Journal of Experimental Botany · doi:10.1093/jxb/err313 · Cited by 555
Lignin plays a negative role in the biochemical process for producing lignocellulosic biofuels
Zeng, Y., Zhao, S., Yang, S., & Ding, S.-Y. (2013) · Current Opinion in Biotechnology · doi:10.1016/j.copbio.2013.09.008 · Cited by 524
Nanda, S., Mohanty, P., Pant, K. K., Naik, S. N., Koziński, J. A., & Dalai, A. K. (2012) · BioEnergy Research · doi:10.1007/s12155-012-9281-4 · Cited by 446
Metadata
- Authors
- Novaes, E., Kirst, M., Chiang, V., Winter-Sederoff, H., & Sederoff, R.
- Affiliations
- North Carolina State University (US)
- School of Forest Resources and Conservation (E.N., M.K.) and University of Florida Genetics Institute (M.K.), University of Florida, Gainesville, Florida 32611; Forest Biotechnology Group, Department of Forestry and Environmental Resources (V.C., R.S.), and Department of Plant Biology (H.W.-S.), North Carolina State University, Raleigh, North Carolina 27695
- Venue
- Plant Physiology
- Publisher
- Oxford University Press (OUP)
- Published
- 2010-10-01
- Volume
- 154
- Issue
- 2
- Pages
- 555-561
- ISSN
- 0032-0889 (ISSN-L)
- 1532-2548
- PMC
- PMC2949025
- OpenAlex
- W2054198220
- Cited by
- 421
- References
- 62
- Citations by year
- 2026: 21 · 2025: 35 · 2024: 36 · 2023: 35 · 2022: 33 · 2021: 40 · 2020: 33 · 2019: 27 · 2018: 31 · 2017: 17 · 2016: 20 · 2015: 15 · 2014: 30 · 2013: 25 · 2012: 13
- FWCI
- 1.26
- Type
- journal article
- Language
- en
- Open access
- bronze
- doi.org
- Licence
- academic.oup.com/journals/pages/open_access/funder_policies/chorus/standard_publication_model (vor)
- Locations
- PLANT PHYSIOLOGY (publishedVersion)
- PubMed Central (publishedVersion)
- PubMed (publishedVersion)
- Funders
- National Science Foundation (0817900)
- U.S. Department of Energy (DE‐FG02‐, DEFG0205ER64114, FG02-05ER64114)
- Consortium for Plant Biotechnology Research
- Office of Science (DEFG0205ER64114)
- Division of Molecular and Cellular Biosciences
- Biological and Environmental Research (DE‐FG02‐05ER64114)
- Topics
- Physical Sciences › Chemical Engineering › Process Chemistry and Technology › Lignin and Wood Chemistry
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Molecular Biology › Plant Gene Expression Analysis
- Physical Sciences › Engineering › Building and Construction › Wood Treatment and Properties
- Keywords
- lignin content, woody biomass, wood formation, negative correlation, monolignol biosynthesis, transcriptional regulation, secondary cell wall, lignin biosynthesis genes, biofuels, paper industry, tree breeding, gene family, metabolic regulation
- MeSH terms
- Lignin, Transcription Factors, Trees, Wood, Gene Expression Regulation, Plant, Biomass
About these checks
Registry, retraction, Cites and statistics results are computed, not judged, with one exception: a printed reference that no index lists is read once by a model before it counts as a citation error. Claims are extracted by a model and kept only when their quote appears verbatim in the text; each cited source is then checked, with a model’s help, for whether it says what the claim relies on. Those verdicts can be wrong — read the claim’s quote and the cited source before relying on one.
Statistics and arithmetic are recomputed, not judged.
Confidence bands are shown only for fields calibrated on our annotated test set. None is calibrated yet, so no band is shown.
What counts as a claim ›How we measure confidence ›How we check this ›
Are you an author, or do you see a problem with this page? Request a review