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Staff Publications

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    'Staff publications' contains references to publications authored by Wageningen University staff from 1976 onward.

    Publications authored by the staff of the Research Institutes are available from 1995 onwards.

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NADP-MALIC ENZYME 1 Affects Germination after Seed Storage in Arabidopsis thaliana
Yazdanpanah, F. ; Maurino, Veronica ; Buijs, G. ; Karimi Jashni, M. ; Willems, L.A.J. ; Sergeeva, L. ; Rajjou, Loïc ; Hilhorst, H.W.M. ; Bentsink, L. - \ 2019
Plant and Cell Physiology 60 (2019)2. - ISSN 0032-0781 - p. 318 - 328.
Aging decreases the quality of seeds and results in agricultural
and economic losses. The damage that occurs at the
biochemical level can alter the seed physiological status.
Although loss of viability has been investigated frequently,
little information exists on the molecular and biochemical
factors involved in seed deterioration and loss of viability.
Oxidative stress has been implicated as a major contributor
to seed deterioration, and several pathways are involved in
protection against this. In this study, we show that seeds of
Arabidopsis thaliana lacking a functional NADP-MALIC
ENZYME 1 (NADP-ME1) have reduced seed viability relative
to the wild type. Seeds of the NADP-ME1 loss-of-function
mutant display higher levels of protein carbonylation than
those of the wild type. NADP-ME1 catalyzes the oxidative
decarboxylation of malate to pyruvate with the simultaneous
production of CO2 and NADPH. Upon seed imbibition,
malate and amino acids accumulate in embryos of aged
seeds of the NADP-ME1 loss-of-function mutant compared
with those of the wild type. NADP-ME1 expression is
increased in imbibed aged as compared with non-aged
seeds. NADP-ME1 activity at testa rupture promotes
normal germination of aged seeds. In seedlings of aged
seeds, NADP-ME1 is specifically active in the root meristematic
zone. We propose that NADP-ME1 activity is required
for protecting seeds against oxidation during seed dry
storage.
Keywords: Ageing Arabidopsis thaliana Enzyme activity
Gene expression Meristematic zone Metabolite
profiling Oxidative stress Seed longevity.
Core features of the hormonal status in in vitro grown potato plants
Kolachevskaya, Oksana O. ; Sergeeva, L. ; Getman, Irina A. ; Lomin, Sergey N. ; Savelieva, E.M. ; Romanov, G.A. - \ 2018
Plant Signaling & Behavior 13 (2018)5. - ISSN 1559-2316
Some time ago, potato transformants expressing Agrobacterium-derived auxin synthesis gene tms1 were generated. These tms1-transgenic plants, showing enhanced productivity, were studied for their hormonal status, turnover and responses in comparison with control plants. For this purpose, contents of phytohormones belonging to six different classes (auxins, cytokinins, gibberellins, abscisic, jasmonic and salicylic acids) were determined by a sensitive UPLC-MS/MS method in tubers and shoots of in vitro grown plants. To date, this study represents the most comprehensive analysis of the potato hormonal system. On the basis of obtained results, several new generalizations concerning potato hormonal status were drawn. Overall, these data can serve as a framework for forthcoming integrative studies of the hormonal system in potato plants.
A disturbed auxin signaling affects adventitious root outgrowth in Solanum dulcamara under complete submergence
Yang, Xinping ; Jansen, Martijn J. ; Zhang, Qian ; Sergeeva, Lidiya ; Ligterink, Wilco ; Mariani, Celestina ; Rieu, Ivo ; Visser, Eric J.W. - \ 2018
Journal of Plant Physiology 224-225 (2018). - ISSN 0176-1617 - p. 11 - 18.
ABA - Adventitious roots - Auxin - Complete submergence - JA - Signaling
Flooding negatively affects the growth and even survival of most terrestrial plants. Upon flooding, the excess water quickly decreases the gas exchange between atmosphere and the submerged plant tissues, which leads to oxygen deficiency resulting in a plant cell energy crisis, and eventually plant death. Solanum dulcamara survives flooding by producing aerenchymatous adventitious roots (ARs) from pre-formed primordia on the stem, which replace the original flood-sensitive root system. However, we found that under complete submergence, AR outgrowth was impaired in S. dulcamara. In the present work, we tried to elucidate the mechanisms behind this phenomenon in particular the involvement of the phytohormones auxin, abscisic acid and jasmonic acid. Abscisic acid (ABA) is a negative regulator of AR outgrowth, but surprisingly the ABA content and signaling were decreased to a similar extent under both partial and complete submergence, suggesting that ABA might not be responsible for the difference in AR outgrowth. Auxin, which is necessary for AR outgrowth, was at similar concentrations in either partially or completely submerged primordia, but complete submergence resulted in a decrease of auxin signaling in the primordia. Application of 1-naphthaleneacetic acid (NAA) to completely submerged plants restored AR outgrowth, implying that auxin response in the rooting tissues of completely submerged plants was reduced. Furthermore, jasmonic acid (JA) concentrations did not differ between partial and complete submergence. To conclude, a disruption in the auxin signaling within S. dulcamara AR primordia may result in the abortion of AR outgrowth under complete submergence.
Natural variation of hormone levels in Arabidopsis roots and correlations with complex root architecture
Lee, Sangseok ; Sergeeva, Lidiya I. ; Vreugdenhil, Dick - \ 2018
Journal of Integrative Plant Biology 60 (2018)4. - ISSN 1672-9072 - p. 292 - 309.
Studies on natural variation are an important tool to unravel the genetic basis of quantitative traits in plants. Despite the significant roles of phytohormones in plant development, including root architecture, hardly any studies have been done to investigate natural variation in endogenous hormone levels in plants. Therefore, in the present study a range of hormones were quantified in root extracts of thirteen Arabidopsis thaliana accessions using a ultra performance liquid chromatography triple quadrupole mass spectrometer. Root system architecture of the set of accessions was quantified, using a new parameter (mature root unit) for complex root systems, and correlated with the phytohormone data. Significant variations in phytohormone levels among the accessions were detected, but were remarkably small, namely less than three-fold difference between extremes. For cytokinins, relatively larger variations were found for ribosides and glucosides, as compared to the free bases. For root phenotyping, length-related traits-lateral root length and total root length-showed larger variations than lateral root number-related ones. For root architecture, antagonistic interactions between hormones, for example, indole-3-acetic acid to trans-zeatin were detected in correlation analysis. These findings provide conclusive evidence for the presence of natural variation in phytohormone levels in Arabidopsis roots, suggesting that quantitative genetic analyses are feasible.
Role of Tulipa gesneriana TEOSINTE BRANCHED1 (TgTB1) in the control of axillary bud outgrowth in bulbs
Moreno-Pachon, Natalia M. ; Mutimawurugo, Marie Chantal ; Heynen, Eveline ; Sergeeva, Lidiya ; Benders, Anne ; Blilou, Ikram ; Hilhorst, Henk W.M. ; Immink, Richard G.H. - \ 2018
Plant Reproduction 31 (2018)2. - ISSN 2194-7953 - p. 145 - 157.
Apical dominance - Axillary bud - Dormancy - TgTB1
Key message: Tulip vegetative reproduction. Abstract: Tulips reproduce asexually by the outgrowth of their axillary meristems located in the axil of each bulb scale. The number of axillary meristems in one bulb is low, and not all of them grow out during the yearly growth cycle of the bulb. Since the degree of axillary bud outgrowth in tulip determines the success of their vegetative propagation, this study aimed at understanding the mechanism controlling the differential axillary bud activity. We used a combined physiological and “bottom-up” molecular approach to shed light on this process and found that first two inner located buds do not seem to experience dormancy during the growth cycle, while mid-located buds enter dormancy by the end of the growing season. Dormancy was assessed by weight increase and TgTB1 expression levels, a conserved TCP transcription factor and well-known master integrator of environmental and endogenous signals influencing axillary meristem outgrowth in plants. We showed that TgTB1 expression in tulip bulbs can be modulated by sucrose, cytokinin and strigolactone, just as it has been reported for other species. However, the limited growth of mid-located buds, even when their TgTB1 expression is downregulated, points at other factors, probably physical, inhibiting their growth. We conclude that the time of axillary bud initiation determines the degree of dormancy and the sink strength of the bud. Thus, development, apical dominance, sink strength, hormonal cross-talk, expression of TgTB1 and other possibly physical but unidentified players, all converge to determine the growth capacity of tulip axillary buds.
Auxin synthesis gene tms1 driven by tuber-specific promoter alters hormonal status of transgenic potato plants and their responses to exogenous phytohormones
Kolachevskaya, Oksana O. ; Sergeeva, Lidia ; Floková, Kristyna ; Getman, Irina A. ; Lomin, Sergey N. ; Alekseeva, Valeriya V. ; Rukavtsova, Elena B. ; Buryanov, Yaroslav I. ; Romanov, Georgy A. - \ 2017
Plant Cell Reports 36 (2017)3. - ISSN 0721-7714 - p. 419 - 435.
Auxin - Cytokinin - Hormonal status - Phytohormones - Potato - Solanum tuberosum - Transformants - Tuberization
Key message: Ectopic auxin overproduction in transgenic potato leads to enhanced productivity accompanied with concerted and occasional changes in hormonal status, and causing altered response of transformants to exogenous auxin or cytokinin.Abstract: Previously, we generated potato transformants expressing Agrobacterium-derived auxin synthesis gene tms1 driven by tuber-specific patatin gene promoter (B33-promoter). Here, we studied the endogenous hormonal status and the response to exogenous phytohormones in tms1 transformants cultured in vitro. Adding indole-3-acetic acid (IAA) or kinetin to culture medium affected differently tuberization of tms1-transformed and control plants, depending also on sucrose content in the medium. Exogenous phytohormones ceased to stimulate the tuber initiation in transformants at high (5–8%) sucrose concentration, while in control plants the stimulation was observed in all experimental settings. Furthermore, exogenous auxin partly inhibited the tuber initiation, and exogenous cytokinin reduced the average tuber weight in most transformants at high sucrose content. The elevated auxin level in tubers of the transformants was accompanied with a decrease in content of cytokinin bases and their ribosides in tubers and most shoots. No concerted changes in contents of abscisic, jasmonic, salicylic acids and gibberellins in tubers were detected. The data on hormonal status indicated that the enhanced productivity of tms1 transformants was due to auxin and not mediated by other phytohormones. In addition, exogenous cytokinin was shown to upregulate the expression of genes encoding orthologs of auxin receptors. Overall, the results showed that tms1 expression and local increase in IAA level in transformants affect both the balance of endogenous cytokinins and the dynamics of tuberization in response to exogenous hormones (auxin, cytokinin), the latter reaction depending also on the carbohydrate supply. We introduce a basic model for the hormonal network controlling tuberization.
Thermodynamic and structural properties of tuber starches from transgenic potato plants grown in vitro and in vivo
Wasserman, L.A. ; Sergeev, A.I. ; Vasil'Ev, V.G. ; Plashchina, I.G. ; Aksenova, N.P. ; Konstantinova, T.N. ; Golyanovskaya, S.A. ; Sergeeva, Lidia ; Romanov, G.A. - \ 2015
Carbohydrate Polymers 125 (2015). - ISSN 0144-8617 - p. 214 - 223.
AtPHYB gene - Crystalline lamellae - DSC - Solanum tuberosum - Starch - Thermodynamic parameters - Transgenic potato - Tubers rol genes

Potato plants harboring Phytochrome B (PHYB) gene from Arabidopsis thaliana or rol genes from Agrobacterium rhizogenes were used to study the effect of transgene expression on structure and properties of starch in tubers. Thermodynamic characteristics of starch (melting temperature, enthalpy of melting, thickness of crystalline lamellae) were shown to be variable depending on the transgene expression and plant culturing mode: in vitro or in soil. The expression of rolB or rolC genes in in vitro cultured plants evoked opposite effects on starch melting temperature and crystalline lamellae thickness. AtPHYB or rolB expression in the soil-grown potato led to the formation of more defective or more ordered starch structures, respectively, in comparison with starches of the same lines grown in vitro. On the whole, our study revealed genotype-dependent differences between starches extracted from tubers of in vitro or in vivo grown plants.

Expression of auxin synthesis gene tms1 under control of tuber-specific promoter enhances potato tuberization in vitro
Kolachevskaya, O.O. ; Alekseeva, V. ; Sergeeva, L. ; Vreugdenhil, D. - \ 2015
Journal of Integrative Plant Biology 57 (2015)9. - ISSN 1672-9072 - p. 734 - 744.
Phytohormones, auxins in particular, play an important role in plant development and productivity. Earlier data showed positive impact of exogenous auxin on potato (Solanum tuberosum L.) tuberization. The aim of this study was to generate potato plants with increased auxin level predominantly in tubers. To this end, a pBinB33-tms1 vector was constructed harboring the Agrobacterium auxin biosynthesis gene tms1 fused to tuber-specific promoter of the class I patatin gene (B33-promoter) of potato. Among numerous independently generated B33:tms1 lines, those without visible differences from control were selected for detailed studies. In the majority of transgenic lines, tms1 gene transcription was detected, mostly in tubers rather than in shoots. Indoleacetic acid (IAA) content in tubers and the auxin tuber-to-shoot ratio were increased in tms1- expressing transformants. The organ-specific increase in auxin synthesis in B33:tms1-transformants accelerated and intensified the process of tuber formation, reduced the dose of carbohydrate supply required for in vitro tuberization, and decreased the photoperiodic dependence of tuber initiation. Overall, a positive correlation was observed between tms1 expression, IAA content in tubers, and stimulation of tuber formation. The revealed properties of B33:tms1 transformants imply an important role for auxin in potato tuberization and offer prospects tomagnify potato productivity by a moderate organ-specific enhancement of auxin content.
Effects of Agrobacterial rol-Genes on the Thermodynamic and Structural Features of Starches Extracted from Potato Microtubers
Wasserman, L.A. ; Aksenova, N.P. ; Konstantinova, T.N. ; Sergeeva, L. ; Golyanovskaya, S.A. ; Krivandin, A.V. ; Romanov, G.A. - \ 2014
Food and Nutrition Sciences 5 (2014)3. - ISSN 2157-944X - p. 250 - 257.
Wild-type potato (Solanum tuberosum L.) plants and their transformants harboring agrobacterial rolB or rolC genes under control of the patatin class I promoter were cultured in vitro. These plants were used as a source of single-node stem cuttings. The structure of native starch in tubers formed on cuttings was determined using methods of X-ray scattering and differential scanning microcalorimetry (DSC). It was found that in starch from tubers of rolB plants the melting temperature of crystalline lamella was lower and their thickness was less than that in wild-type potato. In tubers of rolC plants starch differed from starch in wild-type plants by a higher melting temperature, reduced melting enthalpy, and a greater thickness of crystalline lamellae. The melting of starch from tubers of rolC plants proceeded as the melting of two independent crystalline structures with melting temperatures of 338.0°K and 342.8°K. Overall data show that starches of different structure can be obtained by using transgenic approach
Hormonal Regulation of Tuber Formation in Potato
Aksenova, N.P. ; Sergeeva, L. ; Kolachevskaya, O.O. ; Romanov, G.A. - \ 2013
In: Bulbous Plants: Biotechnology / Ramawat, K.G., Merillon, J-M, New York : CRC Press - ISBN 9781466589674
Tubers of potato (Solanum tuberosum L.) plants serve as organs for vegetative propagation and stock of matter and energy for the next generation. Tuber formation is a complex process involving several successive stages: stolon formation and growth, induction of tuberization, tuber initiation and further growth. Tuber formation is affected by several external and internal factors, fi rst of all photoperiod, temperature, levels of carbohydrates and nitrogen. Adaptive response of tuberization to these cues is a part of the seasonal strategy of potato reproductive development.
Regulation of potato tuber dormancy and sprouting
Aksenova, N.P. ; Sergeeva, L. ; Konstantinova, T.N. ; Golyanovskaya, S.A. ; Kolachevskaya, O.O. ; Romanov, G.A. - \ 2013
Russian Journal of Plant Physiology 60 (2013)3. - ISSN 1021-4437 - p. 301 - 312.
abscisic-acid content - adp-glucose pyrophosphorylase - endogenous aba levels - solanum-tuberosum - gene-expression - in-vitro - postharvest storage - microtuber dormancy - gibberellic-acid - starch synthesis
Dormancy is the final stage of tuber life serving to preserve tubers as organs of vegetative reproduction under unfavorable growth conditions. Since the duration of potato tuber dormancy and their sprouting time have significant economic importance, much attention is given to the study of the regulation of these processes. This review considers metabolite, genetic, and hormonal aspects of regulation of potato (Solanum tuberosum L.) tuber dormancy and sprouting. Particular attention is paid to the relationship between processes occurring in different parts of the tuber: its storage tissues and buds. The interaction of hormonal and metabolite (carbohydrate) regulation of dormancy and sprouting is discussed.
Starch-related Enzymes during Potato Tuber Dormancy and Sprouting
Sergeeva, L.I. ; Claassens, M.M.J. ; Jamar, D.C.L. ; Plas, L.H.W. van der; Vreugdenhil, D. - \ 2012
Russian Journal of Plant Physiology 59 (2012)4. - ISSN 1021-4437 - p. 556 - 564.
adp-glucose pyrophosphorylase - solanum-tuberosum - developmental-changes - hexose-phosphate - gene-expression - metabolism - sucrose - phosphorylase - biosynthesis - tuberization
Activities of enzymes presumably involved in starch biosynthesis (ADP-glucose pyrophosphorylase, AGPase) and/or breakdown (starch phosphorylase, STP; amylases) were determined during potato (Solanum tuberosum L.) tuber dormancy and sprouting. Overall activities of all these enzymes decreased during the first stage of tuber dormancy. No clear changes were detected at the time of dormancy breaking and sprouting. However, when AGPase activity was monitored by in situ staining during the entire dormancy period, a clear decrease during the dormant period and a large increase before visible sprouting could be observed. This increase was especially evident near the vascular tissue and at the apical bud, which showed a very intensive staining. In situ staining of STP activity in sprouting tubers showed that the tissue distribution of STP was the same as for AGPase. As a possible explanation, direct starch cycling is suggested: STP produces glucose-1-phosphate during starch breakdown, which can be directly used as a substrate by AGPase for starch synthesis. Gene expression studies with the AGPaseS promoter coupled to the firefly luciferase reporter gene also clearly showed a higher activity in sprouting tubers as compared to dormant tubers, with the highest expression levels observed around the apical buds. The presence of amylase activity at dormancy initiation and AGPase activity persistent at the sprouting stage suggest that starch was cycling throughout the entire dormancy period. According to the in situ studies, the AGPase activity increased well before visible sprout growth and could therefore be one of the first physiological determinants of dormancy breakage.
Agrobacterial rol genes modify thermodynamic and structural properties of starch in microtubers of transgenic potato
Aksenova, N.P. ; Wasserman, L.A. ; Sergeeva, L.I. ; Konstantinova, T.N. ; Golyanovskaya, S.A. ; Krivandin, A.V. ; Plashchina, I.G. ; Blaszczak, W. ; Fornal, J. ; Romanov, G.A. - \ 2010
Russian Journal of Plant Physiology 57 (2010)5. - ISSN 1021-4437 - p. 656 - 663.
plants - tubers - expression - growth - dna
Wild-type (WT) plants of potato (Solanum tuberosum L.) and their transgenic forms carrying agrobacterial genes rolB or rolC under the control of B33 class I patatin promoter were cultured in vitro on MS medium with 2% sucrose in a controlled-climate chamber at 16-h illumination and 22A degrees C. These plants were used as a source of single-node stem cuttings, which were cultured in darkness on the same medium supplemented with 8% sucrose. The tubers formed on them were used for determination of the structure of native starch using the methods of differential scanning microcalorimetry (DSC), X-ray scattering, and scanning electron microscopy. It was found that, in starch from the tubers of rolB-plants, the temperature of crystalline lamella melting was lower and their thickness was less than in WT potato. In tubers of rolC plants, starch differed from starch in WT plants by a higher melting temperature, considerably reduced melting enthalpy, and a greater thickness of crystalline lamellae. Deconvolution of DSC thermogram makes it possible to interpret the melting of starch from the tubers of rolC plants as the melting of two independent crystalline structures with melting temperatures of 65.0 and 69.8A degrees C. Electron microscopic examination confirmed the earlier obtained data indicating that, in the tubers of rolC plants, starch granules are smaller and in the tubers of rolB plants larger than in WT plants. Possible ways of influence of rol transgenes on structural properties of starch in amyloplasts of potato tubers are discussed
Use of QTL analysis in physiological research
Vreugdenhil, D. ; Koornneef, M. ; Sergeeva, L.I. - \ 2007
Russian Journal of Plant Physiology 54 (2007)1. - ISSN 1021-4437 - p. 10 - 15.
quantitative trait loci - arabidopsis-thaliana - brassica-napus - flowering time - genome - gene - polymorphisms - vernalization - recombinant - metabolism
Quantitative trait locus (QTL) analysis is a powerful approach to map and subsequently identify genes involved in complex traits. Here we describe the basic principles and recent achievements of this method, and its application in physiological research in plants. The rapidly increasing amount of molecular and ¿omics¿ data and genetic resources and tools, in model species (Arabidopsis) and crops, will greatly support and stimulate the use of this approach in the near future.
Genetics of plant performance using Arabidopsis natural variation
Koornneef, M. ; Keurentjes, J.J.B. ; Sergeeva, L.I. ; El-Lithy, M.E.M. ; Barbier, H. ; Ihnatowicz, A. ; Pieper, B. ; Vreugdenhil, D. ; Reymond, M. - \ 2006
Vacuolar invertase regulates elongation of Arabidopsis thaliana roots as revealed by QTL and mutant analysis.
Sergeeva, L.I. ; Keurentjes, J.J.B. ; Bentsink, L. ; Vonk, J. ; Plas, L.H.W. van der; Koornneef, M. ; Vreugdenhil, D. - \ 2006
Proceedings of the National Academy of Sciences of the United States of America 103 (2006)8. - ISSN 0027-8424 - p. 2994 - 2999.
quantitative trait loci - sucrose synthase - genetic-variation - developmental-changes - plant invertases - hexose-phosphate - enzymes - potato - expression - maize
The possible role of the sucrose-splitting enzymes sucrose synthase and invertase in elongating roots and hypocotyls of Arabidopsis was tested by using a combination of histochemical methods and quantitative trait locus (QTL) analysis. Lengths of roots and hypocotyls correlated better with invertase activities than with sucrose synthase activities. The highest correlations were observed with activities in the elongating zones of roots. The genetic basis of these correlations was studied by using QTL analysis. Several loci, affecting invertase activity, colocated with loci that had an effect on root or hypocotyl length. Further fine mapping of a major locus for root length, but not for hypocotyl length (top chromosome 1), consistently showed colocation with the locus for invertase activity containing a gene coding for a vacuolar invertase. The analysis of a functional knockout line confirmed the role of this invertase in root elongation, whereas other invertase genes might play a role in hypocotyl elongation. Thus, we show the power of QTL analysis, combined for morphological and biochemical traits, followed by fine-mapping and mutant analysis, in unraveling the function of genes and their role in growth and development
Integrative approaches for genetical analysis of Arabidopsis thaliana
Keurentjes, J.J.B. ; Sergeeva, L.I. ; Vreugdenhil, D. ; Koornneef, M. - \ 2004
In: Book of Abstracts, Keystone Symposia "Biological Discovery Using Diverse High-Throughput Data Steamboat Springs, Colorado : - p. 46 - 46.
Histochemical analysis reveals organ-specifics Quantitative Trait Loci for enzyme activities in Arabidopsis
Sergeeva, L.I. ; Vonk, J. ; Keurentjes, J.J.B. ; Plas, L.H.W. van der; Koornneef, M. ; Vreugdenhil, D. - \ 2004
Plant Physiology 134 (2004)1. - ISSN 0032-0889 - p. 237 - 245.
wild-type - starchless mutant - qtl analysis - thaliana - phosphoglucomutase - metabolism - deficient - growth - map
To identify genetic loci involved in the regulation of organ-specific enzyme activities, a specific histochemical staining protocol was used in combination with quantitative trait locus (QTL) analysis. Using phosphoglucomutase (PGM) as an example, it is shown that enzyme activity can specifically, and with high resolution, be visualized in non-sectioned seedlings of Arabidopsis. The intensities of staining were converted to quantitative data and used as trait for QTL analysis using Landsberg erecta x Cape Verde Islands recombinant inbred lines. Independently, PGM activities were quantified in whole-seedling extracts, and these data were also used for QTL analysis. On the basis of extract data, six significant (P <0.05) loci affecting PGM activity were found. From the histochemical data, one or more specific QTLs were found for each organ analyzed (cotyledons, shoot apex, hypocotyl, root, root neck, root tip, and root hairs). Loci detected for PGM activity in extracts colocated with loci for histochemical staining. QTLs were found coinciding with positions of (putative) PGM genes but also at other positions, the latter ones supposedly pointing toward regulatory genes. Some of this type of loci were also organ specific. It is concluded that QTL analysis based on histochemical data is feasible and may reveal organ-specific loci involved in the regulation of metabolic pathways.
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