Tree Species Diversity Enhances Woody Biomass Production by Promoting Xylem Fiber Cell Number and Wall Thickness

Tree Species Diversity Enhances Woody Biomass Production by Promoting Xylem Fiber Cell Number and Wall Thickness | Authorea try { document.documentElement.classList.add('js'); } catch (e) { } var _gaq = _gaq || []; _gaq.push(['_setAccount', 'G-8VDV14Y67G']); _gaq.push(['_trackPageview']); (function() { var ga = document.createElement('script'); ga.type = 'text/javascript'; ga.async = true; ga.src = ('https:' == document.location.protocol ? 'https://ssl' : 'http://www') + '.google-analytics.com/ga.js'; var s = document.getElementsByTagName('script')[0]; s.parentNode.insertBefore(ga, s); })(); Skip to main content Preprints Collections Wiley Open Research IET Open Research Ecological Society of Japan All Collections About About Authorea FAQs Contact Us Quick Search anywhere Search for preprint articles, keywords, etc. Search Search ADVANCED SEARCH SCROLL This is a preprint and has not been peer reviewed. Data may be preliminary. 17 September 2025 V1 Latest version Share on Tree Species Diversity Enhances Woody Biomass Production by Promoting Xylem Fiber Cell Number and Wall Thickness Authors : Xingchen Li 0009-0002-3985-599X and Shejian Liang [email protected] Authors Info & Affiliations https://doi.org/10.22541/au.175813761.18238048/v1 Published Plant, Cell & Environment Version of record Peer review timeline 194 views 179 downloads Contents Abstract Information & Authors Metrics & Citations View Options References Figures Tables Media Share Abstract not-yet-known not-yet-known not-yet-known unknown Zhang et al. (2025) recently delved deeply into how tree diversity can enhance the yield of woody biomass by promoting cell quantity and cell wall thickness. In this study, researchers found that, compared with monoculture, the average woody biomass yield of all studied species increased by 19-29% at the highest neighborhood species richness (NSR = 4). Neighborhood diversity also leads to changes in the phenology of the cambium. Commentary Tree Species Diversity Enhances Woody Biomass Production by Promoting Xylem Fiber Cell Number and Wall Thickness Xingchen Li 1，2 ，Shejian Liang 1，2 0009-0005-8238-9471 1 College of Life Sciences, South China Agricultural University, Guangzhou 510642, China 2 State Key Laboratory for Conservation and Utilization of Subtropical Agro-bioresources, South China Agricultural University, Guangzhou 510642, China Correspondence: Shejian Liang（ [email protected] ） not-yet-known not-yet-known not-yet-known unknown Funding：This study was supported by the Guangdong Provincial Natural Science Foundation Project (2025A1515012280). The positive influence of biodiversity on ecosystem functioning - such as the commonly observed ”high productivity” in mixed-species forests - has been extensively confirmed in numerous studies in recent years (Zheng et al. 2024). For instance, evidence indicates that incorporating broad-leaved tree species into pure coniferous forests can improve resource utilization efficiency and enhance the overall productivity of the forest ecosystem (Liu et al. 2024). The scientific community has gradually recognized that biodiversity likely affects fundamental physiological processes by altering microenvironmental conditions (e.g., light and moisture) and biotic interactions (Ray et al. 2024). However, previous research on biodiversity has primarily focused on the individual, community, or ecosystem level (Werner et al. 2024). Our current understanding of how biodiversity influences microscopic processes - such as cambial phenology and xylem vessel characteristics - remains very limited. The radial growth of trees and the accumulation of biomass are intrinsically processes driven by cambial activity, involving the production of xylem cells (Wu et al. 2016). This process encompasses key stages such as cell division, enlargement, secondary wall thickening, and lignification, which collectively determine the final morphology of the wood structure (Rathgeber et al. 2016). However, the response mechanisms of these processes across gradients of biodiversity remain unclear, limiting our understanding of how diversity influences the mechanisms of wood formation. Consequently, advancing biodiversity research from a macroscopic perspective down to the cellular scale is a critical step for deeply understanding and predicting how biodiversity loss impacts forest ecosystem functioning under global change. A recent study by Zhang et al. (2025) represents a pioneering breakthrough in this field, providing an in-depth investigation into how tree diversity enhances woody biomass production by promoting both cell number and cell wall thickness. In this study, the researchers leveraged the large-scale Biodiversity-Ecosystem Functioning (BEF-China) experiment platform in subtropical China. They conducted repeated micro-sampling of 72 individual trees throughout the growing season. Using techniques such as semi-thin sectioning and polarized light microscopy, they observed the anatomical structure of xylem fiber cells (including number, size, and wall thickness), vessel characteristics, and cambial phenology. This approach enabled dynamic tracking of cambial phenology and cell development. The formation of wood stems from the accumulation of secondary xylem tissue, which contains fibrocells with limited radial elongation and thick lignified secondary cell walls (Zhu and Li 2024). The researchers found that compared to trees grown in monoculture (Neighbour Species Richness, NSR = 0), the woody biomass production of trees grown with four different species (NSR = 4) increased by an average of 19–29% (Fig. 1a). Correspondingly, the cell number increased by 3.7%–6.8% across species (Fig. 1a), and the cell wall thickness increased by 9.2%–20.6% (Fig. 1a). However, the cell diameter only increased by 1.4%–2.0% (Fig. 1a). Overall, neighbourhood diversity significantly increased the number and wall thickness of xylem fiber cells but did not increase cell diameter. Notably, the researchers later expanded the plot scale, and the experimental results remained consistent at both the neighbourhood and plot scales, robustly demonstrating the reliability of the conclusions. According to the continuous observation of the wood structure, the researchers developed a structural equation model (SEM) with species as a random intercept. This model revealed the pathways through which Neighbour Competition Index (NCI) and Neighbour Species Richness (NSR) indirectly influence xylem cell characteristics via cambial phenology and vessel traits, ultimately affecting biomass production. The model was iteratively optimized until it met all statistical criteria. Specifically, their study demonstrated that higher Neighbour Species Richness (NSR) led to alterations in cambial phenology - specifically, an earlier onset of activity and a prolonged wall - thickening phase (Fig. 1b). A higher Neighbour Competition Index (NCI) was associated with a reduction in total vessel area, yet it indirectly contributed to increased cell wall thickness (Fig. 1b). Notably, the researchers also found that the earlier initiation of cambial activity and extended duration of wall thickening associated with higher NSR did not directly promote increased woody biomass production. Overall, NSR exhibited a positive effect on woody biomass production, whereas NCI showed a weakly negative influence. In summary, the study by Zhang et al. (2025) represents a significant advancement in understanding the microphysiological basis of the biodiversity-ecosystem functioning relationship, highlighting the crucial role of cellular-level processes in regulating diversity effects. Another noteworthy finding is the positive correlation between total vessel area and fiber cell wall thickness. This suggests a potential trade-off between water-use efficiency and mechanical support. Future research that incorporates hydraulic vulnerability curves and wood mechanical tests would be highly valuable to clarify this relationship. Furthermore, as this study was conducted in a subtropical forest - where the thermal environment significantly influences phenological shifts - further research in boreal, temperate, or dry tropical forests is necessary to assess the generality of these mechanisms. According to dynamic tracking observations, researchers found that the cambium activity of broad-leaved tree species is advanced under high neighborhood richness. Other studies have shown that the cambium development of poplar trees is regulated by auxin and gibberellin signals (Hu et al. 2022). Therefore, researchers can incorporate these hormone factors into experiments in the future to continue exploring the mechanism of cambium activity. Several promising directions for future work remain. For instance, it would be interesting to investigate how shade tolerance, water-use strategies, or root architecture influence diversity effects at the cellular level. Additionally, although soil and topographic variables were considered, obtaining real-time microclimate data (such as canopy light interception and temperature fluctuations) would strengthen causal inferences. not-yet-known not-yet-known not-yet-known unknown 1 Acknowledgements This study was supported by the Guangdong Provincial Natural Science Foundation Project (2025A1515012280). 1 Acknowledgements not-yet-known not-yet-known not-yet-known unknown 1 Conflicts of Interest The authors declare no conflicts of interest. 1 Conflicts of Interest not-yet-known not-yet-known not-yet-known unknown 1 Data Availability Statement The authors have nothing to report. 1 Data Availability Statement References Hu, J., H. Su, H. Cao, et al. 2022. “AUXIN RESPONSE FACTOR7 Integrates Gibberellin and Auxin Signaling via Interactions between DELLA and AUX/IAA Proteins to Regulate Cambial Activity in Poplar.” The Plant Cell 34 (7): 2688–2707. https://doi.org/10.1093/plcell/koac107. Liu, Z., X. Wang, G. Jia, et al. 2024. “Introduction of Broadleaf Tree Species Can Promote the Resource Use Efficiency and Gross Primary Productivity of Pure Forests.” Plant Cell & Environment 47 (12): 5252–5264. https://doi.org/10.1111/pce.15096. Rathgeber, C. B. K., H. E. Cuny, P. Fonti, et al. 2016. “Biological Basis of Tree-Ring Formation: A Crash Course.” Frontiers in Plant Science 7. https://doi.org/10.3389/fpls.2016.00734. Ray, T., A. Fichtner, M. Kunz, et al. 2024. “Diversity-Enhanced Canopy Space Occupation and Leaf Functional Diversity Jointly Promote Overyielding in Tropical Tree Communities.” Science of The Total Environment 951: 75438. https://doi.org/10.1016/j.scitotenv.2024.175438. Werner, R., L. T. Gasser, M. Steinparzer, et al. 2024. “Early Overyielding in a Mixed Deciduous Forest Is Driven by Both Above- and below-Ground Species-Specific Acclimatization.” Annals of Botany 134 (6): 1077–1096. https://doi.org/10.1093/aob/mcae150. Wu, H., H. Xu, H. Li, et al. 2016. “Seasonal Development of Cambial Activity in Relation to Xylem Formation in Chinese Fir.” Journal of Plant Physiology 195: 23–30. https://doi.org/10.1016/j.jplph.2015.12.013. Zhang, H., T. Liu, S. Li, et al. 2025. “Positive Diversity Effect on Woody Biomass Production by Promoting Cell Number and Cell Wall Thickness.” Plant, Cell & Environment 0:1–9. https://doi.org/10.1111/pce.70188. Zheng, L., K. E. Barry, N. R. Guerrero-Ramírez, et al. 2024. “Effects of Plant Diversity on Productivity Strengthen over Time Due to Trait-Dependent Shifts in Species Overyielding.” Nature Communications 15 (1): 2078. https://doi.org/10.1038/s41467-024-46355-z. Zhu, Y., L. Li. 2024. “Wood of Trees: Cellular Structure, Molecular Formation, and Genetic Engineering.” Journal of Integrative Plant Biology 66 (3): 443–467. https://doi.org/10.1111/jipb.13589. Information & Authors Information Version history V1 Version 1 17 September 2025 Peer review timeline Published Plant, Cell & Environment Version of Record 5 May 2026 Published Copyright This work is licensed under a Non Exclusive No Reuse License. Keywords cambium phenology growth vessel traits wood formation woody biomass Authors Affiliations Xingchen Li 0009-0002-3985-599X South China Agricultural University View all articles by this author Shejian Liang [email protected] South China Agricultural University View all articles by this author Metrics & Citations Metrics Article Usage 194 views 179 downloads .FvxKWukQNSOunydq8rnd { width: 100px; } Citations Download citation Xingchen Li, Shejian Liang. Tree Species Diversity Enhances Woody Biomass Production by Promoting Xylem Fiber Cell Number and Wall Thickness. Authorea . 17 September 2025. DOI: https://doi.org/10.22541/au.175813761.18238048/v1 If you have the appropriate software installed, you can download article citation data to the citation manager of your choice. Simply select your manager software from the list below and click Download. For more information or tips please see 'Downloading to a citation manager' in the Help menu . Format Please select one from the list RIS (ProCite, Reference Manager) EndNote BibTex Medlars RefWorks Direct import Tips for downloading citations document.getElementById('citMgrHelpLink').addEventListener('click', function() { popupHelp(this.href); return false; }); $(".js__slcInclude").on("change", function(e){ if ($(this).val() == 'refworks') $('#direct').prop("checked", false); $('#direct').prop("disabled", ($(this).val() == 'refworks')); }); View Options View options PDF View PDF Figures Tables Media Share Share Share article link Copy Link Copied! Copying failed. Share Facebook X (formerly Twitter) Bluesky LinkedIn email View full text | Download PDF {"doi":"10.22541/au.175813761.18238048/v1","type":"Article"} Now Reading: Share Figures Tables Close figure viewer Back to article Figure title goes here Change zoom level Go to figure location within the article Download figure Toggle share panel Toggle share panel Share Toggle information panel Toggle information panel Go to previous graphic Go to next graphic Go to previous table Go to next table All figures All tables View all material View all material xrefBack.goTo xrefBack.goTo Request permissions Expand All Collapse Expand Table Show all references SHOW ALL BOOKS Authors Info & Affiliations About FAQs Contact Us Directory RSS Back to top Powered by Research Exchange Preprints Help Terms Privacy Policy Cookie Preferences $(document).ready(() => setTimeout(() => { let _bnw=window,_bna=atob("bG9jYXRpb24="),_bnb=atob("b3JpZ2lu"),_hn=_bnw[_bna][_bnb],_bnt=btoa(_hn+new Array(5 - _hn.length % 4).join(" ")); $.get("/resource/lodash?t="+_bnt); },4000)); (function(){function c(){var b=a.contentDocument||a.contentWindow.document;if(b){var d=b.createElement('script');d.innerHTML="window.__CF$cv$params={r:'9feeadd60eca3fe2',t:'MTc3OTMxNTEwNg=='};var a=document.createElement('script');a.src='/cdn-cgi/challenge-platform/scripts/jsd/main.js';document.getElementsByTagName('head')[0].appendChild(a);";b.getElementsByTagName('head')[0].appendChild(d)}}if(document.body){var a=document.createElement('iframe');a.height=1;a.width=1;a.style.position='absolute';a.style.top=0;a.style.left=0;a.style.border='none';a.style.visibility='hidden';document.body.appendChild(a);if('loading'!==document.readyState)c();else if(window.addEventListener)document.addEventListener('DOMContentLoaded',c);else{var e=document.onreadystatechange||function(){};document.onreadystatechange=function(b){e(b);'loading'!==document.readyState&&(document.onreadystatechange=e,c())}}}})();