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RIONDATO, ROBERTA (2009) Xilogenesi ed accrescimento longitudinale al limite superiore del bosco: determinanti ambientali e fisiologici. [Tesi di dottorato]

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Abstract (inglese)

During evolution, trees have developed anatomical and structural modifications in order to obtain an adeguate longitudinal growth.
On Earth, some species grow taller than 100 m and a single giant sequoia tree spans about 12 orders of magnitude from seedling to mature tree.
However, vertical growth is not unlimited: every species grows to a maximum which can vary with the different envinronmental conditions.
At present, it is still uncertain which are the actual mechanisms determining the decline in height growth during ontogeny.
Among the scientific community it is commonly accepted the limitations to the height growth are size-related rather than age-related and, particularly, the ability to deliver water from roots to leaves (for the photosynthetic processes) seemed be very important in this respect.
The whole tree architecture must be designed in order to guarantee an efficient long-distance transport and it is determined by mechanical and hydraulic constraints.
Hydraulic constraints arise from well-known physic laws, especially from the Hagen-Poiseuille law.
In fact, the ascent sap flow is affected, in fact, by the path-lenght, the cell radius and the wall roughness that cause high losses of energy and increase the cost of transport (hydraulic resistance).
A number of mechanisms of compensation are known to exist that help to minimize the increment of resistance with path-lenght but great attention have been attribuited to a recent theoretical model (West et al., 1999) that proposed the xylem tapering as a key compensation mechanisms.
The degree of conduit tapering predicted by WBE model rapresents the minimum value on which evolutive forces are converged: a cylindrical pipe, in fact, could induce limitations to height growth.
At present, scientists do not agree either that a minimum and universal treshold of conduit tapering does actually exist and either that this value is invariant during ontogeny.
The aim of this work is to increase the knowledge about the xylem structure, and its ontogenetic changes, in alpine conifers (Picea abies Karst., Larix decidua Mill., Pinus cembra L.) that have approached their maximum height and show physiological top dieback.
Beyond the tapering of conduits, further analyses are carried out in order to search other mechanisms of resistance compensation, , and the effects on xilogenesis of environmental variables are also investigated.
This research was divided in chapters, each dealing with a specific argument.
In the first issue it has been carried out the stem-analysis, a method used to evaluate radial, longitudinal and volumetric growth of the examined trees.
During ontogeny there was a change in the whole stem profile with the stem top, more tapered than the lower part, becoming more cylindrical.
The second and the fourth issues have focused on the hydraulic architecture of three individuals and have evaluated the shape of conduits and their variation with time.
At different heights along the stem, and within each tree-ring, diameter of tracheids has been measured. Woody samples have been extracted along the radial and the longitudinal axis, embedded in paraffin and cut by a rotative microtome. The obtained slides have been observed under a light microscope.
In each tree the longitudinal vessel profile has been analysed for a minimum of 20 years before the top dieback: the results showed a degree of tapering approaching the WBE model prediction.
However, the degree of variation of conduits along the stem was not uniform but rather segmented: at the tree top vessels were more tapered than towards the base. This discrepancy increased when the trees were close to the dieback year and, in such situation, it seems that the hydraulic resistance increased. For this reason, the total number of cells in the cross-sections were measured for different years and heights.
Climate growth relationships have been analyzed in the third chapter, and the results showed that the cell production (ring width) was more related to environmental variables (temperature, precipitation and cloudiness) than cell lumen.
The intensity of responses to climate-signal was different along the tree stem: the most sensible zone was located at the crown base and this could be the cause of changes in cell’s shape and of diameter:height ratio.
This Thesis has introduced a new element to the research about the limits to tree height, in fact it has investigated ontogenetical changes in trees that were approaching their maximum height.
The results, complex and not fully explained, would suggest that an insufficient degree of conduit tapering strongly constrains the height growth of trees.

Abstract (italiano)

Nel corso della loro evoluzione gli alberi hanno sviluppato degli accorgimenti anatomici e strutturali finalizzati ad ottenere un adeguato sviluppo verticale. Sulla Terra non mancano infatti esempi di specie in grado di superare i 100 m di altezza e nel corso della propria esistenza un singolo individuo di sequoia può modificare le sue dimensioni attraversando ben 12 ordini di grandezza.
L’accrescimento longitudinale, tuttavia, non è infinito: ogni specie manifesta una altezza massima e all’interno della stessa si possono osservare dimensioni diverse in relazione all’ambiente di crescita.
I meccanismi che conducono al rallentamento del ritmo di crescita in altezza, fenomeno ontogenetico tipico di tutte le specie vascolari, non sono ancora stati chiariti.
Nella comunità scientifica, a seguito di accesi dibattiti, è sembrata prevalere l’idea che la limitazione sia da imputare alle dimensioni della pianta, piuttosto che alla sua età, ed in particolare alla capacità di trasportare una sufficiente quantità d’acqua dalle radici alle foglie, sedi del processo fotosintetico.
L’intera architettura della pianta sarebbe quindi ideata per ottenere un trasporto efficiente e sarebbe sottoposta a vincoli di natura meccanica e idraulica, questi ultimi descrivibili in base a note leggi della fisica quali quella di Hagen-Poiseuille.
La lunghezza del percorso, la dimensione della cellula xilematica e le asperità delle pareti sono infatti la causa di notevoli perdite energetiche (aumento della resistenza idraulica) che ostacolano il moto ascendente della linfa.
Tra i vari meccanismi di compensazione che la pianta potrebbe adottare per minimizzare l’effetto della lunghezza del percorso un grande interesse ha riscosso quello proposto nel modello WBE (West et al., 1999), ossia la rastremazione xilematica.
È stata perciò ipotizzata l’esistenza di un valore minimo di rastremazione, sul quale tutte le specie sono portate a convergere nel corso dell’evoluzione: un profilo xilematico troppo cilindrico potrebbe infatti indurre una limitazione alla crescita in altezza.
Attualmente non vi è accordo, tra gli studiosi, se esista realmente una soglia di rastremazione minima ed universale e se questa venga mantenuta costante nel corso dell’ontogenesi.
Lo scopo di questo lavoro è proprio quello di indagare la struttura xilematica di conifere di alta quota tipiche delle Alpi (Picea abies Karst., Larix decidua Mill., Pinus cembra L.), valutando i cambiamenti di forma dei condotti di piante giunte ormai alla loro massima altezza e con cimale disseccato per cause fisiologiche.
Inoltre sono state eseguite una serie di analisi collaterali per ricercare eventuali altri meccanismi compensativi che la pianta potrebbe adottare nel tentativo di limitare l’aumento di resistenza idraulica interna allo xilema.
Sono stati valutati anche gli effetti dei principali parametri climatici sulla xilogenesi.
Il lavoro, quindi, si presenta costituito da diversi capitoli, ognuno riguardante uno specifico argomento della ricerca.
Nel primo capitolo, attraverso il metodo dell’analisi del fusto, è stato possibile valutare l’accrescimento radiale, longitudinale e volumetrico delle piante esaminate.
Il profilo del fusto cambia nel corso dell’ontogenesi e la forma del cimale, significativamente diversa da quella delle parti inferiori, diviene sempre più cilindrica.
Nel secondo e quarto capitolo sono state approfondite le conoscenze sull’architettura idraulica di tre soggetti, valutando la forma dei condotti e la sua variazione nel tempo.
Le analisi sono state eseguite misurando il diametro delle tracheidi in ogni anello della pianta, a diverse altezze. I campioni legnosi, estratti lungo gli assi longitudinale e radiale del fusto, sono stati inclusi in paraffina, tagliati con microtomo rotativo e osservati al microscopio.
Per ogni pianta è stato possibile ricostruire il profilo longitudinale di almeno 20 anni precedenti il disseccamento del cimale: ne è risultato un grado di rastremazione prossimo alle soglie previste dal modello WBE. Tuttavia la forma dei condotti non deve ritenersi uniforme ma piuttosto spezzata dato che le cellule nel cimale sono decisamente più rastremate di quelle del fusto. Questa differenza aumenta in prossimità dell’anno del disseccamento suggerendo l’ipotesi che un peggioramento nel sistema di trasporto possa essere la causa di un aumento di resistenza idraulica. A questo proposito è stato stimato anche in numero totale di cellule presente nella sezione trasversale del fusto in vari anni ed a varie altezze.
Nel terzo capitolo si è dato ampio spazio alle relazioni clima-accrescimento ed è stato possibile verificare che il numero di cellule (ovvero lo spessore anulare) è molto più influenzato del lume del condotto dalla tempertura, dalle precipitazioni e dalla nuvolosità.
La pianta, a varie altezze del fusto, mostra risposte diverse: la zona più sensibile si collocherebbe alla base della chioma e questo potrebbe determinare ulteriori variazioni nella forma dei condotti e nei rapporti meccanici tra altezza e diametro.
Questa Tesi contribuisce in maniera innovativa alla ricerca in quanto indaga i cambiamenti ontogenetici in piante già giunte alla loro massima altezza. I risultati, seppur complessi e non completamente interpretabili, contribuiscono a sostenere l’ipotesi che un peggioramento nel sistema di conduzione può pesantemente limitare l’altezza delle piante.

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Tipo di EPrint:Tesi di dottorato
Relatore:COLPI, CRISTIANA
Correlatore:ANFODILLO, TOMMASO
Dottorato (corsi e scuole):Ciclo 21 > Scuole per il 21simo ciclo > TERRITORIO, AMBIENTE, RISORSE E SALUTE > ECOLOGIA
Data di deposito della tesi:28 Gennaio 2009
Anno di Pubblicazione:28 Gennaio 2009
Parole chiave (italiano / inglese):xilogenesi, accrescimento longitudinale, modello WBE, rastremazione xilematica, determinanti ambientali
Settori scientifico-disciplinari MIUR:Area 07 - Scienze agrarie e veterinarie > AGR/05 Assestamento forestale e selvicoltura
Struttura di riferimento:Dipartimenti > Dipartimento Territorio e Sistemi Agro-Forestali
Codice ID:1542
Depositato il:28 Gen 2009
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