Mathesius Group - Root microbe interactions - symbionts to parasites

Legumes form a beneficial symbiosis with nitrogen-fixing bacteria called rhizobia, that leads to the formation of root nodules. We are interested in the molecular mechanisms controlling nodule development in legumes with the long-term aim of expanding nitrogen-fixing associations to reduce nitrogen fertiliser use. We are using a mixture of molecular, cell biological and chemical ecology approaches to understand what features are necessary and unique for nodulation in legumes and what makes the symbiosis functional under biotic stress from soil parasites and pathogens. One focus is on the role of root flavonoids in controlling nodule development and in the communication between roots and soil microbes.

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Selected publications

  • Rae AE, Rolland V, White R, Mathesius U (2021) New methods for confocal imaging of infection threads in crop and model legumes. Plant Methods 7:24.
  • Kawasaki A, Dennis PG, Forstner C, Raghavendra AKH, Mathesius U, Richardson AE, Delhaize E, Gilliham M, Watt M, Ryan PR (2021) Manipulating exudate composition from root apices shapes the microbiome throughout the root system, Plant Physiology, kiab337
  • Qiao Y, Miao S, Jin J, Mathesius U, Tang C (2021) Differential responses of the sunn4 and rdn1-1 super-nodulation mutants of Medicago truncatula to elevated atmospheric CO2. Annals of Botany 128: 441-452
  • Mathesius U, Costa SR (2021) Plant signals differentially affect rhizosphere nematode populations. Journal of Experimental Botany 72: 3496–3499
  • Kawasaki A, Dennis PG, Forstner C, Raghavendra AKH, Richardson AE, Watt M, Mathesius U, Gilliham M,Ryan PR (2021) The microbiome of wheat and rice roots exhibits significant differences in structure between root types and along root axes. Functional Plant Biology, 48: 871-888.
  • Xie Z, Yu Z, Li Y, WangG, Tang C, Mathesius U, Liu X, Liu J, Liu J, Herbert SJ, Wu J, Jin J (2021) Linking rhizospheric diazotrophs to the stimulation of soybean N2 fixation in a Mollisol amended with maize straw. Plant and Soil 463:  279–289.
  • Xie Z, Li Y, Yu Z, Wang G, Tang C,  Mathesius U, Liu X, Liu J, Liu J, Hebert SJ, Wu J, Jin J (2021) Incorporation of maize crop esidue maintains the soybean yield through teh stimulation of nitrogen fixation rather than residue-derived nitrogen in Mollisols.  Field Crops Research 272: 108279.
  • Costa SR, Ng JLP, Mathesius U (2021) Interaction of symbiotic rhizobia and parasitic root-knot nematodes in legume roots – from molecular regulation to field application. Molecular Plant-Microbe Interactions, 34: 470-490.
  • Mens C, Hastwell AH, Su H, Gresshoff PM, Mathesius U, Ferguson BJ. (2020) Characterisation of Medicago truncatula CLE34 and CLE35 in nitrate and rhizobia regulation of nodulation. New Phytologist 229: 2525–2534.
  • Costa, SR, Chin S, Mathesius U (2020). Infection of Medicago truncatula by the root-knot nematode Meloidogyne javanica does not require early nodulation genes. Frontiers in Plant Science 11: 1050
  • Veliz-Vallejos DF, Kawasaki A, Mathesius U (2020) The presence of plant-associated bacteria alters responses to N-acyl homoserine lactone quorum sensing signals that modulate nodulation in Medicago truncatula, Plants, 9: 777
  • Harris JM, Pawlowski K, Mathesius U (2020) Editorial: Evolution of signaling in plant symbioses. Front. Plant Sci. 11: 456.
  • Chapman K, Ivanovici A, Taleski M, Sturrock C, Ng JLP, Mohd-Radzman NA, Frugier F, Bennett M, Mathesius U, Djordjevic MA (2020) CEP receptor signalling controls root system architecture in Arabidopsis and Medicago. New Phytologist 226: 1809–1821.
  • Ng JLP, Welvaert A, Wen J, Chen R, Mathesius U. (2020) The Medicago truncatula PIN2 auxin transporter mediates basipetal auxin transport but is not necessary for nodulation. Journal of Experimental Botany 71: 1562–1573.
  • Demina IV, Maity PJ, Nagchowdhury A, Ng JLP, van der Graaff E, Demchenko KN, Roitsch T, Mathesius U, Pawlowski K (2019) Accumulation of and response to auxins in roots and nodules of the actinorhizal plant Datisca glomerata compared to the model legume Medicago truncatula. Frontiers in Plant Science 10: 1085.
  • Goh C-H, Nicotra AB, Mathesius U (2019) Genes controlling legume nodule numbers affect phenotypic plasticity responses to nitrogen in the presence and absence of rhizobia. Plant, Cell and Environment 42: 1747-1757.
  • Gauthier-Coles C, White R, Mathesius U (2019). Nodulating legumes are distinguished by a sensitivity to cytokinin in the root cortex leading to pseudonodule development. Frontiers in Plant Science 9: 1901.
  • Kawasaki A, Okada S, Zhang C, Delhaize E, Mathesius U, Richardson AE, Watt M, Gilliham M, Ryan PR (2018) A sterile hydroponic system for characterising root exudates from specific root types and whole-root systems of large crop plants. Plant Methods 14: 114.
  • Chin S, Behm CA, Mathesius U (2018) Functions of flavonoids in plant-nematode interactions. Plants 7: 85.
  • Ng JLP, Mathesius U (2018) Acropetal auxin transport inhibition is involved in indeterminate but not determinate nodule formation. Frontiers in Plant Science 9: 169.
  • Kohlen W, Ng JLP, DeinumEE, MathesiusU (2018) Auxin transport, metabolism and signaling during nodule initiation: Indeterminate and determinate nodules. Journal of Experimental Botany 69: 229-244.
  • Li Y, Yu Z, Liu X, Mathesius U, Wang G, Tang C, Wu J, Liu J, Zhang S, Jin J (2017) Elevated CO2 increases nitrogen fixation at the reproductive phase contributing to various yield responses of soybean cultivars. Frontiers in Plant Science 8:1546.
  • Liu Y, Hassan S, Kidd BN, Garg G, Mathesius U, Singh KB, Anderson J (2017) Ethylene signaling is important for isoflavonoid mediated resistance to Rhizoctonia solani in Medicago truncatula. Molecular Plant-Microbe Interactions 30: 691-700.
  • KawasakiA, Donn S, RyanPR, MathesiusU, Devilla R, Jones A, Watt M (2016) Microbiome and exudates of the root and rhizosphere of Brachypodium distachyon, a model for wheat. PLoS ONE 11(10): e0164533.
  • Li YS, Liu XB, Wang GH, Yu ZH,  Mathesius U, Liu JD, Herbert SJ, Jin J (2016) The shift of plant N origins alters C and N assimilation during reproductive stages of soybean grown in a Mollisol. Crop and Pasture Science. 67: 872-880.
  • Van Noorden GE, Verbeek R, Dinh QD, Jin J, Green A, Ng JLP, Mathesius U (2016) Molecular signals controlling the inhibition of nodulation by nitrate in Medicago truncatula. International Journal of Molecular Sciences 17: 1060.
  • Ng JLP, Mathesius U (2016) Measuring auxin transport capacity in seedling roots of Medicago truncatula. Bio-Protocol 6(12): e1842.
  • Ng JLP, Truong TT, Hocart CC, Mathesius U (2016) Quantifying auxin metabolites in young root tissue of Medicago truncatula by liquid chromatography electrospray-ionisation quadrupole time-of-flight (LC-ESI-QTOF) tandem mass spectrometry. Bio-Protocol 6(12): e1843.
  • Schneebeli K, Mathesius U, Zwart AB, Bragg JN, Vogel JP, Watt M (2016) Brachypodium distachyon genotypes vary in resistance to Rhizoctonia solani AG8. Functional Plant Biology 43, 189–198.
  • Goh, C.-H., Nicotra, A.B. and Mathesius U. (2016) The presence of nodules on legume root systems can alter phenotypic plasticity in response to internal nitrogen independent of nitrogen fixation. Plant, Cell and Environment 39: 883-896.
  • Shabala S, White R, Djordjevic MA, Ruan Y-L, Mathesius U (2016) Root to shoot signaling: diverse molecules, pathways and functions. Functional Plant Biology 43: 87-104.
  • Beckmann EA, Estavillo GM, Mathesius U, Djordjevic MA, Nicotra AN (2015) The Plant Detectives: innovative undergraduate teaching to inspire the next generation of plant biologists. Frontiers in Plant Science 6: 729.
  • Ng JLP, Perrine-Walker FM, Wasson AP, Mathesius U (2015) The control of auxin transport in parasitic and symbiotic root–microbe interactions. Plants 4: 606-643.
  • Ng JLP, Hassan, S, Truong TT, Hocart CH, Laffont C, Frugier F, Mathesius U (2015) Flavonoids and auxin transport inhibitors rescue symbiotic nodulation in the Medicago truncatula cytokinin perception mutant cre1. Plant Cell 27: 2210-2226.
  • Schneebeli K, Mathesius U, Watt M (2015) Brachypodium distachyon is a pathosystem model for the study of the wheat root disease Rhizoctonia solani AG 8. Plant Pathology 64: 91-100.
  • Gowland K, Mathesius U, Clements M, Nicotra A (2015) Understanding the distribution of three species of epiphytic orchids in temperate Australian rainforest by investigation of their host and fungal associates. Lankesteriana 7: 1-2.
  • Veliz-Vallejos DF, van Noorden GE, Mengqi Y and Mathesius U (2014) A Sinorhizobium meliloti-specific N-acyl homoserine lactone quorum-sensing signal increases nodule numbers in Medicago truncatula independent of autoregulation. Frontiers in Plant Science 5: 551.
  • George DT, Behm CA , Hall DH, Mathesius U, Rug M, Nguyen KCQ and Verma NK (2014) Shigella flexneri infection in Caenorhabditis elegans: Cytopathological examination and identification of host responses. PLoS ONE 9: e106085.
  • Ferguson BJ and Mathesius U (2014) Phytohormone regulation of legume-rhizobia interactions. Journal of Chemical Ecology 40, 770-790.
  • Kurdyukov S, Mathesius U, Nolan KE, Sheahan MB, Goffard N, Carroll BJ, Rose RJ (2014) The 2HA line of Medicago truncatula has characteristics of an epigenetic mutant that is weakly ethylene insensitive. BMC Plant Biology 14:174.
  • George DT, Mathesius U, Behm CA, Verma NK (2014) The periplasmic enzyme, AnsB, of Shigella flexneri modulates bacterial adherence to host epithelial cells. PLoS ONE 9: e94954.
  • Mortier V, Wasson A, Jaworek P, De Keyser A, Decroos M, Holsters M, Tarkowski P, Mathesius U, Goormachtig S (2014) Role of LONELY GUY genes in indeterminate nodulation on Medicago truncatula. New Phytologist 202: 582–593.
  • Goh C-H, Veliz-Vallejo DF, Nicotra AB, Mathesius U (2013) The impact of beneficial plant-associated microbes on plant phenotypic plasticity. Journal of Chemical Ecology 39:826–839.
  • Cazzonelli, CI, Vanstraelen M, YinK, Carron-ArthurA, Nisar N, Tarle G, Cuttriss AJ, Searle IR, Simon S, Benkova E, Mathesius U, Masle J, Friml J, Pogson BJ (2013) Role of the Arabidopsis PIN6 auxin transporter in auxin homeostasis and auxin-mediated development. PLoS ONE 8: e70069.
  • Weston LA and Mathesius U (2013) Flavonoids: their structure, biosynthesis and role in the rhizosphere, including allelopathy. Journal of Chemical Ecology 39: 283-297.
  • Jin J, Watt M and Mathesius U (2012) The autoregulation gene SUNN mediates changes in root organ formation in response to nitrogen through alteration of shoot-to-root auxin transport. Plant Physiology 159: 489-500.
  • Hassan, S. and Mathesius, U. (2012) The role of flavonoids in root-rhizosphere signaling - opportunities and challenges for improving plant-microbe interactions. Journal of Experimental Botany 63: 3429-3444.
  • Vincent, D., Du Fall, L.A., Livk, A., Mathesius, U., Lipscombe R.J., Oliver, R.P., Freisen, T.L., Solomon, P.S. (2012) A functional genomics approach to dissect the mode-of-action of the Stagonospora nodorum effector protein SnToxA in wheat. Molecular Plant Pathology 13 : 467-482.
  • MathesiusU, Djordjevic MA, Oakes M, Goffard N, Haerizadeh F, Weiller GF, Singh MB, Bhalla PL (2011) Comparative proteomic profiles of the soybean (Glycine max) root apex and differentiated root zone. Proteomics 11, 1707–1719.
  • Teplitski M, Mathesius U and Rumbaugh KB (2011) Perception and degradation of N-acyl homoserine lactone quorum sensing signals by mammalian and plant cells. Chemical Reviews 111: 100-116.
  • Mathesius U and Watt M (2011) Rhizosphere signals for plant-microbe interactions: implications for field-grown plants. In: U. Lüttge et al. (eds.), Progress in Botany 72: 125-161. Springer-Verlag Berlin Heidelberg.
  • Plet J, Wasson A, Ariel F, Le Signor C, Baker D, Mathesius U, Crespi M, and Frugier F (2011) MtCRE1-dependent cytokinin signaling integrates bacterial and plant cues to coordinate symbiotic nodule organogenesis in Medicago truncatula. Plant Journal 65, 622–633.
  • Ignjatovic V, Lai C, Summerhayes R, Mathesius U, Tawfilis S, Perugini MA, Monagle P (2011) Age-related differences in plasma proteins: How plasma proteins change from neonates to adults. PLoS ONE 6: e17213
  • Nicotra AB, Atkin, OK, Bonser SP, Davidson,A, Finnegan EJ, Mathesius U, Poot P, Purugganan MD, Richards CL, Valladares F, van Kleunen M (2010) Plant phenotypic plasticity in a changing climate. Trends in Plant Science 15: 684-692.
  • Hassan S, Behm CA and Mathesius U (2010) Effectors of plant parasitic nematodes that re-program root cell development. Functional Plant Biology 37: 933–942.
  • Laffont C, Blanchet S, Lapierre C, Brocard L, Ratet P, Crespi M, Mathesius U and Frugier F (2010) The Compact Root Architecture 1 gene regulates lignification, flavonoid production and polar auxin transport in Medicago truncatula. Plant Physiology 153:1597–1607.
  • Mathesius U (2010) The role of auxin in root-symbiont and root-pathogen interactions – from development to defense. Progress in Botany 71: 185-210. (Eds. Lüttge, U.E., Beyschlag, W., Büdel, B., Francis, D. ). Springer Verlag Berlin, Heidelberg.
  • Grunewald W., van Noorden G.E., van Isterdael G., Beeckman T., Gheysen G. and Mathesius U (2009). Manipulation of auxin transport in plant roots during Rhizobium symbiosis and nematode parasitism. Plant Cell 21: 2553-2562.
  • Shane MW, Ngo H, Lambers H, Pate JS, Cawthray GR, Mathesius U, Canny MJ, McCully ME (2009) Summer dormancy and winter growth: root survival strategy in a perennial monocotyledon. New Phytologist. 183: 1085-1096
  • Wasson AP, Ramsay K, Jones MGK and Mathesius U (2009) Differing requirements for flavonoids during the formation of lateral roots, nodules and root knot nematode galls in Medicago truncatula. New Phytologist 183: 167–179
  • Mathesius U (2009) Comparative proteomics studies of root-microbe interactions. J Proteomics 72: 353-366.
  • Mathesius U (2008). Auxin – at the root of nodule development? Functional Plant Biology, 35: 651-668.
  • Oelkers K, Goffard N, Weiller FG, Gresshoff PM, Mathesius U, Frickey T (2008) Bioinformatic analysis of the CLE signaling peptide family. BMC Plant Biology 8:1.
  • van Noorden GE, Kerim T, Goffard N, Wiblin R, Pellerone FI, Rolfe BG, Mathesius U (2007) Overlap of proteome changes in Medicago truncatula in response to auxin and Sinorhizobium meliloti. Plant Physiology 144: 1115-1131.
  • de JongF, MathesiusU, Imin N, Rolfe BG. (2007) A proteome study of the proliferation of cultured Medicago truncatula protoplasts. Proteomics 7: 722-736.
  • Beveridge C, Mathesius U, Rose RJ, Gresshoff PM (2007) Common regulatory themes in meristem development and whole plant homeostasis. Current Opinion in Plant Biology 10: 44-51.
  • Prayitno J, Imin N., Rolfe B.G., Mathesius U. (2006) Identification of ethylene-mediated protein changes during nodulation in Medicago truncatula using proteome analysis. Journal of Proteome Research, 5: 3084-3095.
  • Prayitno J, Rolfe B.G., Mathesius U. (2006) The ethylene insensitive sickle mutant of Medicago truncatula shows altered auxin transport regulation during nodulation. Plant Physiology, 142: 168-180.
  • Förster, B., Mathesius U. and Pogson B.P. (2006) Comparative proteomics of high light stress in the model alga Chlamydomonas reinhardtii. Proteomics, 6: 4309-4320.
  • Wasson, A.P., Pellerone, F.I. and Mathesius U. (2006) Silencing the flavonoid pathway in Medicago truncatula inhibits root nodule formation and prevents auxin transport regulation by rhizobia. Plant Cell 18, 1617-1629.
  • van Noorden, G.E., Ross, J.J, Reid, J.B., Rolfe, B.G. and U. Mathesius (2006) Defective long distance auxin transport regulation in the Medicago truncatula super numeric nodulation mutant. Plant Physiology 140: 1494-1506.
  • Bauer W.D., Teplitski, M., Mathesius, U. (2005) Plants deal with bacterial quorum sensing. ASM News 71: 129-135.
  • Bauer, W. D. and Mathesius, U. (2004) Plant responses to bacterial quorum sensing signals. Current Opinion in Plant Biology 7: 429-433
  • Imin, N., de Jong, F., Mathesius, U., van Noorden, G., Saeed, N., Wang, X-D., Rose, R.J., and Rolfe, B.G (2004) Proteome reference maps of Medicago truncatula embryogenic cell cultures generated from single protoplasts. Proteomics 4: 1883-1896
  • Ferguson, B.F. and Mathesius U. (2003) Signaling interactions during nodule development.  Journal of Plant Growth Regulation, 22: 47-72.
  • Mathesius, U. (2003) Conservation and divergence of signalling pathways between roots and soil microbes - the Rhizobium-legume symbiosis compared to the development of lateral roots, mycorrhizal interactions and nematode-induced galls. Plant and Soil 255: 105-119.
  • Rolfe, B.G., Mathesius U., Djordjevic, M., Weinman, .J, Hocart, C., Weiller G., and Bauer, W.D. (2003) Proteomic analysis of legume-microbe interactions Comparative and Functional Genomics 4: 225-228. (IF 0.3) (Ci 11)
  • Mathesius U., Mulders, S., Gao, M., Teplitski, M., Caetano-Anolles, G., Rolfe, B. G. and Bauer, W. D. (2003) Extensive and specific responses of a eukaryote to bacterial quorum sensing signals. Proceedings of the National Academy of Science USA 100: 1444-1449.
  • Mathesius, U., Imin, N., Chen, H., Djordjevic, M.A., Weinman, J.J., Natera, S.H.A., Morris, A.C., Kerim, T., Paul, S., Menzel, C., Weiller, G.F., and Rolfe, B.G. (2002) Evaluation of proteome reference maps for cross-species identification of proteins by peptide mass fingerprinting. Proteomics 2: 1288-1303.
  • Mathesius, U., Keijzers, G., Natera, S. H. A., Weinman, J. J., Djordjevic, M. A. and Rolfe, B. G. (2001) Establishment of a root proteome reference map for the model legume Medicago truncatula using the EST database for peptide mass fingerprinting. Proteomics, 1: 1424-1440.
  • Mathesius, U. (2001). Flavonoids induced in cells undergoing nodule organogenesis in white clover are regulators of auxin breakdown by peroxidase. Journal of Experimental Botany 52: 419-426. 
  • Mistrik, I., Pavlovkin, J., Wächter, R., Pradel, K. S., Schwalm, K., Hartung, W., Mathesius, U., Stöhr, C., and Ullrich, C. I. (2000). Impact of Agrobacterium tumefaciens-induced stem tumors on NO3- uptake in Ricinus communis.  Plant and Soil 226: 87-98.
  • Mathesius, U., Charon, C., Rolfe, B. G., Kondorosi, A., and Crespi, M. (2000). Temporal and spatial order of events during the induction of cortical cell divisions in white clover by Rhizobium inoculation or localised cytokinin addition. Molecular Plant-Microbe Interactions 13: 617-628.
  • Mathesius, U., Weinman, J. J. Rolfe, B. G., and Djordjevic, M. A. (2000).  Rhizobia can induce nodules in white clover by “hijacking” mature cortical cells activated during lateral root development. Molecular Plant-Microbe Interactions 13: 170-182.
  • Hutangura, P., Mathesius, U., Rolfe, B. G. and Jones, M. E. K. (1999). Auxin induction is a trigger for root gall formation caused by root-knot nematodes in white clover and is associated with the activation of the flavonoid pathway. Australian Journal of Plant Physiology 26: 221-231.
  • Müller, A. and Mathesius, U. (1999). The paleoenvironments of coastal lagoons in the southern baltic sea I. The application of sedimentay Corg/N ratios as source indicators of organic matter. Paleogeography, Paleoclimatology, Paleoecology 145: 1-16.
  • Mathesius, U, Schlaman, H. R. M., Spaink, H. P., Sautter, C., Rolfe, B. G. and Djordjevic, M. A. (1998).  Auxin transport inhibition precedes nodule formation in white clover roots and is regulated by flavonoids and derivatives of chitin oligosaccharides. Plant Journal 14: 23-34.
  • Mathesius, U., Bayliss, C., Weinman, J. J., Schlaman, H. R. M., Spaink, H. P., Rolfe, B. G., McCully, M. E., and Djordjevic, M. A. (1998). Flavonoids synthesised in cortical cells during nodule initiation are early developmental markers in white clover. Molecular Plant-Microbe Interactions 11: 1223-1232.
  • Rolfe, B. G., Djordjevic, M. A., Weinman, J. J., Mathesius, U., Pittock, C., Gärtner, E., Ride, K. M., Dong, Z., McCully, M. E,. and Mc Iver, J. (1997). Root morphogenesis in legumes and cereals and the effect of bacterial inoculation on root development. Plant and Soil 194: 131-144.
  • Djordjevic, M. A., Mathesius, U., Arioli, T., Weinman, J. J., and Gärtner, E. (1997). Chalcone synthase gene expression in transgenic subterranean clover correlates with localised accumulation of flavonoids. Australian Journal of Plant Physiology 24: 119-132.
  • Larkin, P. J., Gibson, J. M., Mathesius, U, Weinman, J. J., Gärtner, E., Hall, E., Tanner, G. J., Rolfe, B. G. and Djordjevic, M. A. (1996). Transgenic white clover. Studies with the auxin responsive promoter, GH3, in root gravitropism and lateral root development. Transgenic Research 5: 325-335.
  • Cleary, A. L. and Mathesius, U. (1996). Rearrangements of F-actin during stomatogenesis visualised by confocal microscopy in fixed and permeabilised Treadescantia leaf epidermis. Botanica Acta 109: 15-24.

All publications

Zooming in on legumes offers hope for nitrogen pollution crisis

Story | Tuesday 2 November 2021
While Angus Rae focuses intently on a captivating microscopic world, he is actually working towards a solution for an environmental catastrophe occurring on a global scale.

Nodulation in legumes

Story | Thursday 13 April 2017
Legumes are an interesting plant to study due to their symbiotic relationship with nitrogen fixing bacteria called Rhizobia, which are housed within specialised root structures called nodules. The work of ANU researchers has been very important for our understanding of symbiosis, nodule formation and nitrogen fixation.

ARC Discovery projects and DECRA fellowships

Story | Monday 17 November 2014

ARC grant success

Story | Thursday 3 November 2011

Innovative teaching recognised

Story | Friday 1 July 2011
Recorded Seminar (Ulrike Mathesius)

RSB Director's Seminar: Are legumes different? Origins and consequences of evolving nitrogen fixing symbioses

Event | Mon 15 April 2024
Professor Ulrike Mathesius, Head of Division, Division of Plant Sciences

PS Seminar Series: Plant Ca2+-electrical signaling - Light-gate channels optogenetics break new grounds

Event | Wed 6 March 2024
Cytosolic Ca2+ signals and changes in pH are universal signaling elements that couple a wide range of stimuli to their characteristic responses in plants. Despite decades of intensive research, it is still poorly understood.

PS Seminar Series - Building Eco-Resilience of Oil Palm with Biotechnology Tools towards a Green Planet

Event | Wed 21 February 2024
Since 2000, oil palm cultivation has generated considerable controversy: more than 20 million ha of plantations have been linked with deforestation, burning, a high carbon footprint, biodiversity loss and environmental pollution from the palm oil industry.