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1.
Modelling seasonal dynamics of secondary growth in R
Jernej Jevšenak, Jožica Gričar, Sergio Rossi, Peter Prislan, 2022, original scientific article

Abstract: The monitoring of seasonal radial growth of woody plants addresses the ultimate question of when, how and why trees grow. Assessing the growth dynamics is important to quantify the effect of environmental drivers and understand how woody species will deal with the ongoing climatic changes. One of the crucial steps in the analyses of seasonal radial growth is to model the dynamics of xylem and phloem formation based on increment measurements on samples taken at relatively short intervals during the growing season. The most common approach is the use of the Gompertz equation, while other approaches, such as general additive models (GAMs) and generalised linear models (GLMs), have also been tested in recent years. For the first time, we explored artificial neural networks with Bayesian regularisation algorithm (BRNNs) and show that this method is easy to use, resistant to overfitting, tends to yield s-shaped curves and is therefore suitable for deriving temporal dynamics of secondary tree growth. We propose two data processing algorithms that allow more flexible fits. The main result of our work is the XPSgrowth() function implemented in the radial Tree Growth (rTG) R package, that can be used to evaluate and compare three modelling approaches: BRNN, GAM and the Gompertz function. The newly developed function, tested on intra-seasonal xylem and phloem formation data, has potential applications in many ecological and environmental disciplines where growth is expressed as a function of time. Different approaches were evaluated in terms of prediction error, while fitted curves were visually compared to derive their main characteristics. Our results suggest that there is no single best fitting method, therefore we recommend testing different fitting methods and selection of the optimal one.
Keywords: artificial neural networks, cambium, generalized additive model, Gompertz function, growing season, intra-annual time series
Published in DiRROS: 21.07.2022; Views: 463; Downloads: 312
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2.
Transition dates from earlywood to latewood and early phloem to late phloem in Norway Spruce
Jožica Gričar, Katarina Čufar, Klemen Eler, Vladimir Gryc, Hanuš Vavrčík, Martin De Luis, Peter Prislan, 2021, original scientific article

Abstract: Climate change will affect radial growth patterns of trees, which will result in different forest productivity, wood properties, and timber quality. While many studies have been published on xylem phenology and anatomy lately, little is known about the phenology of earlywood and latewood formation, also in relation to cambial phenology. Even less information is available for phloem. Here, we examined year-to-year variability of the transition dates from earlywood to latewood and from early phloem to late phloem in Norway spruce (Picea abies) from three temperate sites, two in Slovenia and one in the Czech Republic. Data on xylem and phloem formation were collected during 2009-2011. Sensitivity analysis was performed to determine the specific contribution of growth rate and duration on wood and phloem production, separately for early and late formed parts. We found significant differences in the transition date from earlywood to latewood between the selected sites, but not between growth seasons in trees from the same site. It occurred in the first week of July at PAN and MEN and more than two weeks later at RAJ. The duration of earlywood formation was longer than that of latewood formation; from 31.4 days at PAN to 61.3 days at RAJ. In phloem, we found differences in transition date from early phloem to late phloem also between the analysed growth seasons; from 2.5 weeks at PAN to 4 weeks at RAJ Compared to the transition from earlywood to latewood the transition from early phloem to late phloem occurred 25-64 days earlier. There was no significant relationship between the onset of cambial cell production and the transition dates. The findings are important to better understand the inter-annual variability of these phenological events in spruce from three contrasting temperate sites, and how it is reflected in xylem and phloem anatomy.
Keywords: Picea abies, xylem formation, phloem formation, cambium, tracheid, sieve cell, conifer, temperate environment
Published in DiRROS: 22.03.2021; Views: 1097; Downloads: 808
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3.
Timeline of autumn phenology in temperate deciduous trees
Inge Dox, Jožica Gričar, Lorène Julia Marchand, Sebastien Leys, Paolo Zuccarini, Charly Geron, Peter Prislan, Bertold Mariën, Patrick Fonti, Holger Lange, Josep Peñuelas, Jan Van den Bulcke, Matteo Campioli, 2020, original scientific article

Abstract: Cessation of xylem formation or wood growth (CWG) and onset of foliar senescence (OFS) are key autumn phenological events in temperate deciduous trees. Their timing is fundamental for development and survival of trees, ecosystem nutrient cycling, the seasonal exchange of matter and energy between the biosphere and atmosphere and affect the impact and feedback of forests to global change. A large-scale experimental effort and improved observational methods have allowed us to compare the timing of CWG and OFS for different deciduous tree species in Western Europe, in particularly silver birch, a pioneer species, and European beech, a late-succession species, at stands of different latitudes, of different levels of site fertility, and for two years with contrasting meteorological and drought conditions i.e., the low-moderately dry 2017 and the extremely dry 2018. Specifically, we tested whether foliar senescence started before, after or concurrently with CWG. OFS and CWG occurred generally between late September and early November, with larger differences across species and sites for OFS. Foliar senescence started concurrently with CWG in most cases, except for the drier 2018 and, for beech, at the coldest site, where OFS occurred significantly later than CWG. Behavior of beech in Spain, the southern edge of its European distribution, was unclear, with no CWG, but very low wood growth at the time of OFS. Our study suggests that OFS is generally triggered by the same drivers of CWG or when wood growth decreases in late summer, indicating an overarching mechanism of sink limitation as a possible regulator of the timing of foliar senescence.
Keywords: autumn phenology, xylem formation, foliar senescence, cambium, chlorophyll, radial growth, wood, decidiuous trees, common aspen, common beech, pedunculate oak, silver birch
Published in DiRROS: 10.06.2020; Views: 1873; Downloads: 1121
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4.
Post-fire effects on development of leaves and secondary vascular tissues in Quercus pubescens
Jožica Gričar, Polona Hafner, Martina Lavrič, Mitja Ferlan, Nives Ogrinc, Bor Krajnc, Klemen Eler, Dominik Vodnik, 2020, original scientific article

Abstract: An increased frequency of fire events on the Slovenian Karst is in line with future climate-change scenarios for drought-prone environments worldwide. It is therefore of the utmost importance to better understand tree-fire-climate interactions for predicting the impact of changing environment on tree functioning. To this purpose, we studied the post-fire effects on leaf development, leaf carbon isotope composition (%13C), radial growth patterns and the xylem and phloem anatomy in undamaged (H-trees) and fire-damaged trees (F-trees) of Q. pubescens with good re-sprouting ability in spring 2017, the growing season after a rangeland fire in August 2016. We found that the fully developed canopy of F-trees reached only half of the LAI values measured in H-trees. Throughout the season, F-trees were characterised by higher water potential and stomatal conductivity and achieved higher photosynthetic rates compared to unburnt H-trees. The foliage of F-trees had more negative %13C values than those of H-trees. This reflects that F-trees less frequently meet stomatal limitations due to reduced transpirational area and more favourable leaf-to-root ratio. In addition, the growth of leaves in F-trees relied more on the recent photosynthates than on reserves due to the fire disturbed starch accumulation in the previous season. Cambial production stopped 3 weeks later in F-trees, resulting in 60% and 22% wider xylem and phloem increments, respectively. A novel approach by including phloem anatomy in the analyses revealed that fire caused changes in conduit dimensions in the early phloem but not in the earlywood. However, premature formation of the tyloses in the earlywood vessels of the youngest two xylem increments in F-trees implies that xylem hydraulic integrity was also affected by heat. Analyses of secondary tissues showed that although xylem and phloem tissues are interlinked changes in their transport systems due to heat damage are not necessarily coordinated.
Keywords: pubescent oak, cambium, radial growth, xylem, phloem, anatomy, sub-Mediterranean
Published in DiRROS: 19.03.2020; Views: 2378; Downloads: 1270
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5.
Seasonal radial growth of Black pine (Pinus nigra Arnold) from Bosnia and Herzegovina, monitored by the pinning method and manual band dendrometers
Simon Poljanšek, Jernej Jevšenak, Jožica Gričar, Tom Levanič, 2019, original scientific article

Abstract: Despite numerous dendroclimatological investigations into different tree species from Bosnia and Herzegovina, information is lacking on intra-annual wood formation patterns, which would help us to interpret the climate signal in tree rings better. Using the pinning method and manual band dendrometers, we investigated the seasonal dynamics of radial growth of black pine (Pinus nigra Arnold) trees in two successive growing seasons: 2011 and 2012. The up to 60-year-old trees grew in a stand at the base of a hill in the western, mountainous part of the Balkan Peninsula. The seasonal dynamics of wood formation and final number of cells differed between the studied years. Wood formation started in both years in early to mid-March. Differences were noticed in the wood production culmination; in 2011 it occurred at the end of May and beginning of June in 2012 and 2011, respectively. Xylem growth finished in 2012 in the middle of August and in 2011 in the middle of September. Based on the first derivative of the Gompertz function calculated rate of xylem growth was lowest in 2011. The dendrometers recorded a slow increment rate in spring, higher in summer and a decreasing rate again in the late summer in both growing seasons. In comparison with pinning, dendrometers showed a delay in the start of radial growth of up to 20 days in 2012. Additionally, dendrometers showed an increase in stem girth after the end of both growing seasons, when wood formation was already completed. Deviations between the two methods could be ascribed to the influence of water storage dynamics in the main stem and numerous structural processes in bark tissue, which are captured in dendrometer data. The influence of weather conditions on xylem phenology is also indicated by differences between the two studied years, although it is difficult to identify the influence of particular short-term weather events.
Keywords: pinning, manual dendrometers, radial growth, Pinus nigra, cambium, Balkan Peninsula
Published in DiRROS: 11.09.2019; Views: 4629; Downloads: 3835
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6.
Odziv kambija navadne smreke (Picea abies) na ogrevanje in hlajenje debla
Jožica Gričar, Martin Zupančič, Katarina Čufar, Primož Oven, 2004, original scientific article

Abstract: Proučili smo vpliv eksperimentalnega ogrevanja in hlajenja dela debla navadne smreke (Picea abies) na kambijevo aktivnost in celično diferenciacijo. Poskusasta potekala 30 dni; ogrevanje od 29.3.2004 do 3.5.2004, hlajenje pa od14.6.2004 do 20.7.2004. Vzorce floema, kambija in ksilema smo iz dreves odvzeli vsakih 10 dni, pripravili prečne prereze tkiv in preparate opazovali ssvetlobnim mikroskopom. Lokalno ogrevanje je po 10 dneh induciralo delitveno aktivnost kambija na floemsko stran, po 20 dneh pa tudi na ksilemsko. Po 30 dneh je pri ogrevanem vzorcu nastalo do 15 celic ranega lesa. V tem času se jeredna delitvena aktivnost kambija pri kontrolnem drevesu šele začela. Odziv kambija na hlajenje debla je bil manj izrazit. Anatomskih razlik med kontrolnim in hlajenim vzorcem po 10 in 20 dneh ni bilo. Po 30 dneh je pri hlajenem vzorcu začel nastajati kasni les, pri kontrolnem drevesu pa šele prehodni rani-kasni les. S poskusom smo demonstrirali, da je mogoče pri smrekiz umetnim ogrevanjem in hlajenjem debla vplivati na ksilo- in floemogenezo.
Keywords: navadna smreka, Picea abies, kambij, celična diferenciacija, ksilem, floem, ogrevanje, hlajenje, svetlobna mikroskopija, Norway spruce, cambium, cell differentiation, xylem, phloem, heating, cooling, light microscopy
Published in DiRROS: 12.07.2017; Views: 4531; Downloads: 2006
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7.
Annual cambial rhythm in Pinus halepensis and Pinus sylvestris as indicator for climate adaptation
Peter Prislan, Jožica Gričar, Martin De Luis, Klemen Novak, Edurne Martinez Del Castillo, Uwe Schmitt, Gerald Koch, Jasna Štrus, Polona Mrak, Magda Tušek-Žnidarič, Katarina Čufar, 2016, original scientific article

Abstract: To understand better the adaptation strategies of intra-annual radial growth in Pinus halepensis and Pinus sylvestris to local environmental conditions, we examined the seasonal rhythm of cambial activity and cell differentiation at tissue and cellular levels. Two contrasting sites differing in temperature and amount of precipitation were selected for each species, one typical for their growth and the other represented border climatic conditions, where the two species coexisted. Mature P. halepensis trees from Mediterranean (Spain) and sub-Mediterranean (Slovenia) sites, and P. sylvestris from sub-Mediterranean (Slovenia) and temperate (Slovenia) sites were selected. Repeated sampling was performed throughout the year and samples were prepared for examination with light and transmission electron microscopes. We hypothesized that cambial rhythm in trees growing at the sub-Mediterranean site where the two species co-exist will be similar as at typical sites for their growth. Cambium in P. halepensis at the Mediterranean site was active throughout the year and was never truly dormant, whereas at the sub-Mediterranean site it appeared to be dormant during the winter months. In contrast, cambium in P. sylvestris was clearly dormant at both sub-Mediterranean and temperate sites, although the dormant period seemed to be significantly longer at the temperate site. Thus, the hypothesis was only partly confirmed. Different cambial and cell differentiation rhythms of the two species at the site where both species co-exist and typical sites for their growth indicate their high but different adaptation strategies in terms of adjustment of radial growth to environmental heterogeneity, crucial for long-term tree performance and survival.
Keywords: Aleppo pine, Pinus halepensis, Scots pine, Pinus sylvestris, cambium, light microscopy, Mediterranean environment, temporate environment, transmission electron microscopy, xylem
Published in DiRROS: 12.07.2017; Views: 3077; Downloads: 1404
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