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  1. Broughton WJ, Hoh CH, Behm CA, Tung HF
    Planta, 1978 Jan;139(2):183-92.
    PMID: 24414160 DOI: 10.1007/BF00387146
    The sequence of events leading up to the establishment of symbiotic nitrogen-fixation were studied in two tropical legumes, Centrosema pubescens Benth, and Vigna unguiculata L. Walp. Parameters measured included fresh and dry weights, chlorophyll and leghaemoglobin contents, as well as the activities of NADH-nitrate reductase (EC 1.6.6.1), and nitrogenase (nitric-oxide reductase-EC 1.7.99.2) in plants that were inoculated with suitable rhizobia or which were watered with potassium nitrate. Dry weight and photosynthetic activity of both species followed the sigmoidal pattern which is characteristic of most plants. Growth was little different in either a qualitative or quantitative sense whether nitrogen was supplied as nitrate or through dinitrogen fixation. Although the biochemical sequence of events was dependent on the limiting sensitivities of the individual assays used, the data suggest that nitrate reductase is the first measurable enzymatic activity in the nodules (and roots), followed by acetylene reduction and leghaemoglobin in that order. It is possible therefore, that low levels of symbiotic nitrogen fixation occur in the nodules in the absence of leghaemoglobin. Nitrate reductase activity in C. pubescens nodules was negatively exponentially correlated with nitrogenase activity of the same nodules, suggesting a changing metabolism in old nodules. These data are discussed in terms of environmental and physical factors known to control nitrogen fixation.
  2. Wicaksono A, Mursidawati S, Sukamto LA, Teixeira da Silva JA
    Planta, 2016 Aug;244(2):289-96.
    PMID: 27059028 DOI: 10.1007/s00425-016-2512-8
    MAIN CONCLUSION: The propagation of Rafflesia spp. is considered to be important for future development of ornamental and other applications. Thus far, the only successful propagation technique has been grafting. This mini-review succinctly emphasizes what is known about Rafflesia species. Members of the genus Rafflesia (Rafflesiaceae), which are holoparasitic plants known to grow on a host vine, Tetrastigma sp., are widely spread from the Malayan Peninsula to various islands throughout Indonesia. The plant's geographical distribution as well as many other aspects pertaining to the basic biology of this genus have still not been studied. The young flower buds and flowers of wild Rafflesia hasseltii Suringar, Rafflesia keithii Meijer and Rafflesia cantleyi Solms-Laubach are used in local (Malaysia and Indonesia) traditional ethnomedicine as wound-healing agents, but currently no formal published research exists to validate this property. To maintain a balance between its ethnomedicinal and ornamental use, and conservation, Rafflesia spp. must be artificially cultivated to prevent overexploitation. A successful method of vegetative propagation is by host grafting using Rafflesia-impregnated Tetrastigma onto the stem of a normal Tetrastigma plant. Due to difficulties with culture contamination in vitro, callus induction was only accomplished in 2010 for the first time when picloram and 2,4-D were added to a basal Murashige and Skoog medium, and the tissue culture of holoparasitic plants continues to be extremely difficult. Seeds harvested from fertile fruit may serve as a possible method to propagate Rafflesia spp. This paper provides a brief synthesis on what is known about research related to Rafflesia spp. The objective is to further stimulate researchers to examine, through rigorous scientific discovery, the mechanisms underlying the ethnomedicinal properties, the flowering mechanisms, and suitable in vitro regeneration protocols that would allow for the fortification of germplasm conservation.
  3. Ngoot-Chin T, Zulkifli MA, van de Weg E, Zaki NM, Serdari NM, Mustaffa S, et al.
    Planta, 2021 Feb 05;253(2):63.
    PMID: 33544231 DOI: 10.1007/s00425-021-03567-7
    MAIN CONCLUSION: Karyotyping using high-density genome-wide SNP markers identified various chromosomal aberrations in oil palm (Elaeis guineensis Jacq.) with supporting evidence from the 2C DNA content measurements (determined using FCM) and chromosome counts. Oil palm produces a quarter of the world's total vegetable oil. In line with its global importance, an initiative to sequence the oil palm genome was carried out successfully, producing huge amounts of sequence information, allowing SNP discovery. High-capacity SNP genotyping platforms have been widely used for marker-trait association studies in oil palm. Besides genotyping, a SNP array is also an attractive tool for understanding aberrations in chromosome inheritance. Exploiting this, the present study utilized chromosome-wide SNP allelic distributions to determine the ploidy composition of over 1,000 oil palms from a commercial F1 family, including 197 derived from twin-embryo seeds. Our method consisted of an inspection of the allelic intensity ratio using SNP markers. For palms with a shifted or abnormal distribution ratio, the SNP allelic frequencies were plotted along the pseudo-chromosomes. This method proved to be efficient in identifying whole genome duplication (triploids) and aneuploidy. We also detected several loss of heterozygosity regions which may indicate small chromosomal deletions and/or inheritance of identical by descent regions from both parents. The SNP analysis was validated by flow cytometry and chromosome counts. The triploids were all derived from twin-embryo seeds. This is the first report on the efficiency and reliability of SNP array data for karyotyping oil palm chromosomes, as an alternative to the conventional cytogenetic technique. Information on the ploidy composition and chromosomal structural variation can help to better understand the genetic makeup of samples and lead to a more robust interpretation of the genomic data in marker-trait association analyses.
  4. Mazumdar P, Singh P, Kethiravan D, Ramathani I, Ramakrishnan N
    Planta, 2021 May 08;253(6):119.
    PMID: 33963935 DOI: 10.1007/s00425-021-03636-x
    MAIN CONCLUSION: This review provides insights into the molecular interactions between Phytophthora infestans and tomato and highlights research gaps that need further attention. Late blight in tomato is caused by the oomycota hemibiotroph Phytophthora infestans, and this disease represents a global threat to tomato farming. The pathogen is cumbersome to control because of its fast-evolving nature, ability to overcome host resistance and inefficient natural resistance obtained from the available tomato germplasm. To achieve successful control over this pathogen, the molecular pathogenicity of P. infestans and key points of vulnerability in the host plant immune system must be understood. This review primarily focuses on efforts to better understand the molecular interaction between host pathogens from both perspectives, as well as the resistance genes, metabolomic changes, quantitative trait loci with potential for improvement in disease resistance and host genome manipulation via transgenic approaches, and it further identifies research gaps and provides suggestions for future research priorities.
  5. Muniandy K, Tan MH, Shehnaz S, Song BK, Ayub Q, Rahman S
    Planta, 2020 Feb 01;251(2):57.
    PMID: 32008119 DOI: 10.1007/s00425-020-03349-7
    MAIN CONCLUSION: The rice leaf mitochondrial DNA is  more methylated compared to the rice grain mitochondrial DNA. The old rice leaf mitochondrial DNA has also a higher methylation level than the young rice leaf mitochondrial DNA. The presence of DNA methylation in rice organelles has not been well characterized. We have previously shown that cytosine methylation of chloroplast DNA is different between leaf and grain, and varies between young and old leaves in rice. However, the variation in cytosine methylation of mitochondrial DNA is still poorly characterized. In this study, we have investigated cytosine methylation of mitochondrial DNA in the rice grain and leaf. Based on CpG, CHG, and CHH methylation analyses, the leaf mitochondrial DNA was found to be  more methylated compared to the grain mitochondrial DNA. The methylation of the leaf mitochondrial DNA was also higher in old compared to young leaves. Differences in methylation were observed at different cytosine positions of the mitochondrial DNA between grain and leaf, although there were also positions with a similar level of high methylation in all the tissues examined. The differentially methylated cytosine positions in rice mitochondrial DNA were observed mostly in the intergenic region and in some mitochondrial-specific genes involved in ATP production, transcription, and translation. The functional importance of cytosine methylation in the life cycle of rice mitochondria is still to be determined.
  6. Raghavan V
    Planta, 1968 Mar;81(1):38-48.
    PMID: 24519595 DOI: 10.1007/BF00385513
    The metabolism of RNA and protein in the gametophytes of bracken fern (Pteridium aquilinum) is affected by the quality of light in which they are grown. When sporelings were grown as two-dimensional gametophytes in blue light, particulate fractions separated from the sporelings exhibited greater incorporation of uridine-(3)H and leucine-(3)H into RNA and protein, respectively, than those from sporelings grown as one-dimensional protonema in red light. After various periods of exposure of gametophytes to red or blue light in the presence of uridine-(3)H, the nuclei-rich fraction showed the highest specific activity in RNA, and irrespective of incubation time, blue light was more effective than red light. The possibility that enhanced synthesis of RNA in the nucleus in response to blue light is significantly related to the morphological growth pattern of the gametophytes, is discussed.
  7. Mabhaudhi T, Chimonyo VGP, Hlahla S, Massawe F, Mayes S, Nhamo L, et al.
    Planta, 2019 Sep;250(3):695-708.
    PMID: 30868238 DOI: 10.1007/s00425-019-03129-y
    Orphan crops can contribute to building resilience of marginal cropping systems as a climate chnage adaptation strategy. Orphan crops play an important role in global food and nutrition security, and may have potential to contribute to sustainable food systems under climate change. Owing to reports of their potential under water scarcity, there is an argument to promote them to sustainably address challenges such as increasing drought and water scarcity, food and nutrition insecurity, environmental degradation, and employment creation under climate change. We conducted a scoping review using online databases to identify the prospects of orphan crops to contribute to (1) sustainable and healthy food systems, (2) genetic resources for future crop improvement, and (3) improving agricultural sustainability under climate change. The review found that, as a product of generations of landrace agriculture, several orphan crops are nutritious, resilient, and adapted to niche marginal agricultural environments. Including such orphan crops in the existing monocultural cropping systems could support more sustainable, nutritious, and diverse food systems in marginalised agricultural environments. Orphan crops also represent a broad gene pool for future crop improvement. The reduction in arable land due to climate change offers opportunities to expand the area under their production. Their suitability to marginal niche and low-input environments offers opportunities for low greenhouse gas (GHG) emissions from an agro-ecosystems, production, and processing perspective. This, together with their status as a sub-set of agro-biodiversity, offers opportunities to address socio-economic and environmental challenges under climate change. With research and development, and policy to support them, orphan crops could play an important role in climate-change adaptation, especially in the global south.
  8. Tanzi AS, Eagleton GE, Ho WK, Wong QN, Mayes S, Massawe F
    Planta, 2019 Sep;250(3):911-931.
    PMID: 30911885 DOI: 10.1007/s00425-019-03141-2
    MAIN CONCLUSION: Winged bean is popularly known as "One Species Supermarket" for its nutrient-dense green pods, immature seeds, tubers, leaves, and mature seeds. This underutilised crop has potential beneficial traits related to its biological nitrogen-fixation to support low-input farming. Drawing from past knowledge, and based on current technologies, we propose a roadmap for research and development of winged bean for sustainable food systems. Reliance on a handful of "major" crops has led to decreased diversity in crop species, agricultural systems and human diets. To reverse this trend, we need to encourage the greater use of minor, "orphan", underutilised species. These could contribute to an increase in crop diversity within agricultural systems, to improve human diets, and to support more sustainable and resilient food production systems. Among these underutilised species, winged bean (Psophocarpus tetragonolobus) has long been proposed as a crop for expanded use particularly in the humid tropics. It is an herbaceous perennial legume of equatorial environments and has been identified as a rich source of protein, with most parts of the plant being edible when appropriately prepared. However, to date, limited progress in structured improvement programmes has restricted the expansion of winged bean beyond its traditional confines. In this paper, we discuss the reasons for this and recommend approaches for better use of its genetic resources and related Psophocarpus species in developing improved varieties. We review studies on the growth, phenology, nodulation and nitrogen-fixation activity, breeding programmes, and molecular analyses. We then discuss prospects for the crop based on the greater understanding that these studies have provided and considering modern plant-breeding technologies and approaches. We propose a more targeted and structured research approach to fulfil the potential of winged bean to contribute to food security.
  9. Omidvar V, Abdullah SN, Izadfard A, Ho CL, Mahmood M
    Planta, 2010 Sep;232(4):925-36.
    PMID: 20635097 DOI: 10.1007/s00425-010-1220-z
    The 1,053-bp promoter of the oil palm metallothionein gene (so-called MSP1) and its 5' deletions were fused to the GUS reporter gene, and analysed in transiently transformed oil palm tissues. The full length promoter showed sevenfold higher activity in the mesocarp than in leaves and 1.5-fold more activity than the CaMV35S promoter in the mesocarp. The 1,053-bp region containing the 5' untranslated region (UTR) gave the highest activity in the mesocarp, while the 148-bp region was required for minimal promoter activity. Two positive regulatory regions were identified at nucleotides (nt) -953 to -619 and -420 to -256 regions. Fine-tune deletion of the -619 to -420 nt region led to the identification of a 21-bp negative regulatory sequence in the -598 to -577 nt region, which is involved in mesocarp-specific expression. Gel mobility shift assay revealed a strong interaction of the leaf nuclear extract with the 21-bp region. An AGTTAGG core-sequence within this region was identified as a novel negative regulatory element controlling fruit-specificity of the MSP1 promoter. Abscisic acid (ABA) and copper (Cu(2+)) induced the activity of the promoter and its 5' deletions more effectively than methyl jasmonate (MeJa) and ethylene. In the mesocarp, the full length promoter showed stronger inducibility in response to ABA and Cu(2+) than its 5' deletions, while in leaves, the -420 nt fragment was the most inducible by ABA and Cu(2+). These results suggest that the MSP1 promoter and its regulatory regions are potentially useful for engineering fruit-specific and inducible gene expression in oil palm.
  10. Abd-Hamid NA, Ahmad-Fauzi MI, Zainal Z, Ismail I
    Planta, 2020 Feb 18;251(3):68.
    PMID: 32072251 DOI: 10.1007/s00425-020-03356-8
    The SCF complex is a widely studied multi-subunit ring E3 ubiquitin ligase that tags targeted proteins with ubiquitin for protein degradation by the ubiquitin 26S-proteasome system (UPS). The UPS is an important system that generally keeps cellular events tightly regulated by purging misfolded or damaged proteins and selectively degrading important regulatory proteins. The specificity of this post-translational regulation is controlled by F-box proteins (FBPs) via selective recognition of a protein-protein interaction motif at the C-terminal domain. Hence, FBPs are pivotal proteins in determining the plant response in multiple scenarios. It is not surprising that the FBP family is one of the largest protein families in the plant kingdom. In this review, the roles of FBPs, specifically in plants, are compiled to provide insights into their involvement in secondary metabolites, plant stresses, phytohormone signalling, plant developmental processes and miRNA biogenesis.
  11. Gerszberg A
    Planta, 2018 Nov;248(5):1037-1048.
    PMID: 30066219 DOI: 10.1007/s00425-018-2961-3
    The main goal of this publication is an overview of the biotechnological achievements concerning in vitro cultures and transformation of Brassica oleracea var. capitata. Faced with the requirements of the global food market, intensified work on the genetic transformation of economically important plants is carried out in laboratories around the world. The development of efficient procedures for their regeneration and transformation could be a good solution for obtaining, in a shorter time than by traditional methods, plants with desirable traits. Furthermore, conventional breeding methods are insufficient for crop genetic improvement not only because of being time-consuming but also because they are severely limited by sexual incompatibility barriers. This problem has been overcome by genetic engineering, which seems to be a very good technique for cabbage improvement. Despite the huge progress that has been made in the field of plant regeneration and transformation methods, up to now, no routine transformation procedure has been developed in the case of cabbage. This problem stems from the fact that the efficiency of cabbage transformation is closely related to the genotype and some varieties are recalcitrant to transformation. It is obvious that it is not possible to establish one universal regeneration and transformation protocol for all varieties of cabbage. Therefore, it seems fully justified to develop the above-mentioned procedures for individual economically important cultivars. Despite the obstacles of cabbage transformation in laboratories of many countries, especially those where this vegetable is extremely popular (e.g., China, India, Korea, Malaysia, Pakistan), such attempts are made. This article reviews the achievements in the field of tissue culture and cabbage transformation from the last two decades.
  12. Tökés ZA, Woon WC, Chambers SM
    Planta, 1974 Mar;119(1):39-46.
    PMID: 24442407 DOI: 10.1007/BF00390820
    At least two proteases are present in the secretion of the pitchers of Nepenthes macferlanei, a major one with an estimated molecular weight of 59000 and a minor one of 21000. The specificity of the major enzyme, nepenthesin, was broader than previously reported, and strikingly similar to that of pepsin. Lipase activity was also demonstrated, while no amylase activity was present.
  13. Wooi KC, Broughton WJ
    Planta, 1979 Jan;145(5):487-95.
    PMID: 24317866 DOI: 10.1007/BF00380104
    Axenic cultures of bacteroid-containing protoplasts were isolated from root nodules of Vigna unguiculata L. Walp. Dimensions of the protoplasts were 35 to 135 μm long x 35 to 95 μm wide. Yields were about 30 to 50 mg dry weight per gram fresh weight of nodules. About 5x10(8) protoplasts packed into 1 ml of basal medium under the influence of gravity. When incubated in hypertonic, nitrogen-free media, freshly isolated protoplasts began to reduce acetylene to ethylene after a lag period of 24 to 48 h. Various additions to the basal medium showed that the system possessed functional glycolytic and tricarboxylic acid pathways. Endogenous application of various intermediary metabolites stimulated both acetylene reduction and respiration, though not often equally. As acetylene reduction, but not respiration, was inhibitable by both asparagine and glutamine, the system appears suitable for the study of mechanisms controlling symbiotic nitrogen fixation.
  14. Singh P, Mazumdar P, Harikrishna JA, Babu S
    Planta, 2019 Nov;250(5):1387-1407.
    PMID: 31346804 DOI: 10.1007/s00425-019-03246-8
    MAIN CONCLUSION: Rice sheath blight research should prioritise optimising biological control approaches, identification of resistance gene mechanisms and application in genetic improvement and smart farming for early disease detection. Rice sheath blight, caused by Rhizoctonia solani AG1-1A, is one of the most devasting diseases of the crop. To move forward with effective crop protection against sheath blight, it is important to review the published information related to pathogenicity and disease management and to determine areas of research that require deeper study. While progress has been made in the identification of pathogenesis-related genes both in rice and in the pathogen, the mechanisms remain unclear. Research related to disease management practices has addressed the use of agronomic practices, chemical control, biological control and genetic improvement: Optimising nitrogen fertiliser use in conjunction with plant spacing can reduce spread of infection while smart agriculture technologies such as crop monitoring with Unmanned Aerial Systems assist in early detection and management of sheath blight disease. Replacing older fungicides with natural fungicides and use of biological agents can provide effective sheath blight control, also minimising environmental impact. Genetic approaches that show promise for the control of sheath blight include treatment with exogenous dsRNA to silence pathogen gene expression, genome editing to develop rice lines with lower susceptibility to sheath blight and development of transgenic rice lines overexpressing or silencing pathogenesis related genes. The main challenges that were identified for effective crop protection against sheath blight are the adaptive flexibility of the pathogen, lack of resistant rice varieties, abscence of single resistance genes for use in breeding and low access of farmers to awareness programmes for optimal management practices.
  15. Gregory PJ, Mayes S, Hui CH, Jahanshiri E, Julkifle A, Kuppusamy G, et al.
    Planta, 2019 Sep;250(3):979-988.
    PMID: 31250097 DOI: 10.1007/s00425-019-03179-2
    MAIN CONCLUSION: Crops For the Future (CFF), as an entity, has established a broad range of research activities to promote the improvement and adoption of currently underutilised crops. This paper summarises selected research activities at Crops For the Future (CFF) in pursuit of its mission 'to develop solutions for diversifying future agriculture using underutilised crops'. CFF is a research company focussed on the improvement of underutilised crops, so that they might be grown and consumed more widely with benefits to human food and nutritional security; its founding guarantors were the Government of Malaysia and the University of Nottingham. From its base in Malaysia, it engages in research around the world with a focus on species and system diversification. CFF has adopted a food system approach that adds value by delivering prototype food, feed and knowledge products. Bambara groundnut (Vigna subterranea) was adopted as an exemplar crop around which to develop CFF's food system approach with emphasis on the short-day photoperiod requirement for pod-filling and the hard-to-cook trait. Selective breeding has allowed the development of lines that are less susceptible to photoperiod but also provided a range of tools and approaches that are now being exploited in other crops such as winged bean (Psophocarpus tetragonolobus), amaranth (Amaranthus spp.), moringa (Moringa oleifera) and proso (Panicum miliaceum) and foxtail (Setaria italica) millets. CFF has developed and tested new food products and demonstrated that several crops can be used as feed for black soldier fly which can, in turn, be used to feed fish thereby reducing the need for fishmeal. Information about underutilised crops is widely dispersed; so, a major effort has been made to develop a knowledge base that can be interrogated and used to answer practical questions about potential exploitation of plant and nutritional characteristics. Future research will build on the success with Bambara groundnut and include topics such as urban agriculture, rural development and diversification, and the development of novel foods.
  16. Mayes S, Ho WK, Chai HH, Gao X, Kundy AC, Mateva KI, et al.
    Planta, 2019 Sep;250(3):803-820.
    PMID: 31267230 DOI: 10.1007/s00425-019-03191-6
    MAIN CONCLUSION: Bambara groundnut has the potential to be used to contribute more the climate change ready agriculture. The requirement for nitrogen fixing, stress tolerant legumes is clear, particularly in low input agriculture. However, ensuring that existing negative traits are tackled and demand is stimulated through the development of markets and products still represents a challenge to making greater use of this legume. World agriculture is currently based on very limited numbers of crops, representing a significant risk to food supplies, particularly in the face of climate change which is expected to increase the frequency of extreme events. Minor and underutilised crops can help to develop a more resilient and nutritionally dense future agriculture. Bambara groundnut [Vigna subterranea (L.) Verdc.[, as a drought resistant, nitrogen-fixing, legume has a role to play. However, as with most underutilised crops, there are significant gaps in knowledge and also negative traits such as 'hard-to-cook' and 'photoperiod sensitivity to pod filling' associated with the crop which future breeding programmes and processing methods need to tackle, to allow it to make a significant contribution to the well-being of future generations. The current review assesses these factors and also considers what are the next steps towards realising the potential of this crop.
  17. Wong JH, Namasivayam P, Abdullah MP
    Planta, 2012 Feb;235(2):267-77.
    PMID: 21874349 DOI: 10.1007/s00425-011-1506-9
    Phenylalanine ammonia lyase (PAL) plays a major role in plant growth, development and adaptation. In Arabidopsis thaliana, the enzyme is encoded by four genes, namely PAL1, PAL2, PAL3, and PAL4 with PAL1 and PAL2 being closely related phylogenetically and functionally. PAL1 promoter activities are associated with plant development and are inducible by various stress agents. However, PAL2 promoter activities have not been functionally analysed. Here, we show that the PAL2 promoter activities are associated with the structural development of a plant and its organs. This function was inducible in an organ-specific manner by the avirulent strain of Pseudomonas syringae pv. tomato (JL1065). The PAL2 promoter was active throughout the course of the plant development particularly in the root, rosette leaf, and inflorescence stem that provide the plant with structural support. In aerial organs, the levels of PAL2 promoter activities were negatively correlated with relative positions of the organs to the rosette leaves. The promoter was inducible in the root following an inoculation by JL1065 in the leaf suggesting PAL2 to be part of an induced defence system. Our results demonstrate how the PAL2 promoter activities are being coordinated and synchronised for the structural development of the plant and its organs based on the developmental programme. Under certain stress conditions the activity may be induced in favour of certain organs.
  18. Omar FN, Hafid HS, Samsu Baharuddin A, Mohammed MAP, Abdullah J
    Planta, 2017 Sep;246(3):567-577.
    PMID: 28620814 DOI: 10.1007/s00425-017-2717-5
    MAIN CONCLUSION: X-ray microtomography results revealed that delignification process damaged the oil palm fibers, which correlated well with reduction of lignin components and increase of the phenolic content. Biodegradation investigation of natural fibers normally focuses on physico-chemical analysis, with less emphasis on physical aspect like fiber structures affect from microbial activity. In this work, the performance of Pycnoporus sanguineus to delignify oil palm empty fruit bunch fibers through solid-state fermentation utilizing various ratio of POME sludge was reported. In addition to tensile testing, physico-chemical and X-ray microtomography (µ-CT) analyses on the oil palm fibers were conducted to determine the effectiveness of the degradation process. The best ratio of fiber to fungi (60:40) was chosen based on the highest lignin loss and total phenolic content values and further investigation was performed to obtain fermentation kinetics data of both laccase and manganese peroxidase. µ-CT results revealed that delignification process damaged the pre-treated and untreated fibers structure, as evident from volume reduction after degradation process. This is correlated with reduction of lignin component and increase of the phenolic content, as well as lower stress-strain curves of the pre-treated fibers compared to the untreated ones (from tensile testing). It is suggested that P. sanguineus preferred to consume the outer layer of the fiber, before it penetrates through the cellular structure of the inner fiber.
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