Growing on the Edge Hydraulic Architecture of Mangroves Ecological Plasticity and Functional Significance of Water Conducting Tissue in Rhizophora Mucronata and Avicennia Marina 1st Edition Nele Schmitz download pdf
Growing on the Edge Hydraulic Architecture of Mangroves Ecological Plasticity and Functional Significance of Water Conducting Tissue in Rhizophora Mucronata and Avicennia Marina 1st Edition Nele Schmitz download pdf
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Growing on the edge
Print: DCL Print & Sign, Zelzate
All rights reserved. No parts of this book may be reproduced or transmitted in any form or by
any means, electronic, mechanical, photocopying, recording, or otherwise, without the prior
written permission of the author and the publisher.
Growing on the edge
Hydraulic architecture of mangroves:
ecological plasticity and functional significance
of water conducting tissue in Rhizophora mucronata
and Avicennia marina
Thesis
by
Nele Schmitz
May 2008
Promotors
Nico Koedam
Laboratory of Plant Biology and Nature Management
Vrije Universiteit Brussel
Hans Beeckman
Laboratory of Wood Biology and Xylarium
Royal Museum for Central Africa
This research project was a collaboration between the Laboratory of Plant Biology and Nature Management
of the Vrije Universiteit Brussel and the Laboratory of Wood Biology and Xylarium of the Royal Museum
for Central Africa, Tervuren and was funded by the Institute for the Promotion of Innovation through Science
and Technology in Flanders. Research stays abroad were sponsored by the European Commission‘s Research
Infrastructure Action via the Synthesys Project and by the European Cooperation in the field of Scientific
and Technical Research programme (COST) on Cell wall macromolecules and reaction wood (CEMARE)
via a Short Term Scientific Mission. Fieldwork expeditions were financially supported by the National Fund
for Scientific Research (FWO, Belgium), the Schure-Beijerinck-Popping Fonds (Koninklijke Nederlandse
Akademie van Wetenschappen, The Netherlands) and the Flemish Interuniversity Council (VLIR).
General introduction 11
Summarizing Abstract 25
Summarizing Abstract 65
References 101
Samenvatting 123
Acknowledgements 127
8
Outline of the thesis
Wood anatomy can be studied in view of the technical applications of wood, tree pathology, ecology,
dendrochronology but also hydraulics. In this work, the introduction addresses the CHALLENGE for
mangrove trees to transport water in their hydraulically stressful environment and the wood anatomical
APPROACH to understand the way mangroves successfully respond to these high environmental
demands. The current knowledge about the water transport of trees at risk of bubble formation is
presented together with the possible solutions to minimize or to prevent these negative impacts.
However, despite their ultimate position to study the regulation of the water transport under stress
conditions, insight in the hydraulic structure of mangroves, its ecological plasticity and functional
significance remains extremely scanty. The HYPOTHESES tested in this study are presented. The first
part of the thesis highlights the hydraulic architecture of mangroves in relation to the specific
requirements of the regularly flooded and saline environment. The study focuses on two species,
Rhizophora mucronata and Avicennia marina, growing under contrasting conditions but with a certain
overlap. The FIRST CHAPTER starts with the influence of soil water salinity on vessel characteristics
and ends with the question why vessel diameters are nearly constant. The SECOND CHAPTER aims to
answer this question by searching for a trend in intervessel pit anatomy. The remarkable variation in
intervessel pit membrane thickness and electron density is the subject of the THIRD CHAPTER, presenting
a study of the topochemistry of the intervessel pits of both mangrove species. The second part
of the thesis focuses on the formation of wood by successive cambia and its implications for the
hydraulic architecture. While the anatomical development of growth layers via successive cambia has
been described for some species, the periodicity of the growth layer formation as a whole has never
been addressed before. In the FOURTH CHAPTER the annual character of the growth layers of Avicennia
marina is investigated while CHAPTER FIVE proposes a growth mechanism via successive cambia and
the potential ecological and functional advantage of this peculiar anatomical structure for life in the
mangrove environment. The thesis is completed with a general conclusion on the plasticity of the
water transport system of mangrove trees, the functional advantage of having successive cambia and
this all in view of their survival in an environment characterized by exceptional growth conditions,
which are also to some extent unpredictable.
9
General Introduction
10
General Introduction
G eneral
Introduction
Structure
11
General Introduction
From the origin of life on land, water uptake and transport against gravitational forces and under
the restrictions imposed by the environment are one of the major challenges for plants (Kramer and
Boyer, 1995; Kozlowski and Pallardy, 1997; Maseda and Fernández, 2006). Both for growth and
species distribution water availability clearly is one of the most determining factors for land plants
(e.g. Ball, 1998; Brodribb and Hill, 1999; Pockman and Sperry, 2000; Maherali et al., 2004). This is
especially true for tall growing trees since tree height is constrained by water-controlled physiological
factors (Woodward, 2004; Zaehle, 2005; Holbrook and Zwieniecki, 2008). Nevertheless, a successful
forest ecosystem with trees reaching heights of several tens of meters developed along the coasts
of the tropical and subtropical climate zone: mangroves (Tomlinson, 1994; Spalding et al., 1997;
Kathiresan and Bingham, 2001). Spontaneously the question arises how mangrove trees manage
to survive under these sometimes extreme but almost permanently limiting conditions for water
transport. More specifically, what is the effect of the salty water and the alternation between periods
of drought and periods of flooding on mangrove functioning, which is to a large extent the transport of
water? To understand tree hydraulics the primary need is to understand the structure of the hydraulic
network, which will at once clarify species distributions and limits on tree height (e.g. Tyree and
Ewers, 1991; Hacke et al., 2001a; Anfodillo et al., 2006; Loepfe et al., 2007; Petit et al., 2007; Sperry
et al., 2008).
This study wishes to contribute to unravel the hydraulic structure of mangrove trees with the aim to
gain insight in its ecological plasticity and functionality. First, clarification for two basic questions
must be sought: (1) “What makes the mangrove environment so stressful for transporting water?”
and (2) “How can ecological wood anatomy help in understanding mangrove functioning under the
stressful environmental conditions of the mangrove habitat?”.
12
General Introduction
dynamic because of the daily but also the monthly rhythm (e.g. Hoguane et al., 1999; Bala Krishna
Prasad and Ramanathan, 2005). A fluctuating salinity brings more suffering to plants than a constant
salinity of the same mean (Lin and Sternberg, 1993). The degree of fluctuation in pore water salinity
of the substrate depends on tidal amplitude, the position along the coastal slope, as well as on local
hydrological characteristics (Kitheka, 1996, 1997, 1998) that can be affected by human action (Moritz-
Zimmermann et al., 2002; Gopal and Chauhan, 2005; Liu, 2005).
The mangrove forest of Gazi Bay in Kenya, the study area, has a tidal amplitude of about 3.8 m with
a maximum of 4.1 m (Kenya Ports Authority tide tables for Kilindini, Mombasa) and is characterized
by a sloping topography (Matthijs et al., 1999). The resulting gradient in tidal elevation parallels
the zoned species distribution. Two major species of the forest in Gazi are Rhizophora mucronata
Lam., restricted to the seaward side of the forest with the exception of the upstream area of the river
Kidogoweni where salinity is influenced by fresh river water, and Avicennia marina (Forssk.) Vierh.
with a disjunct distribution in the forest, occurring both at the seaside and the landward side where
trees are stunted (Beeckman et al., 1990; Dahdouh-Guebas et al., 2004). For most other mangrove
species, the highly dynamic conditions of the landward zone are unbearable (Clough, 1984; Naidoo,
1985). Due to its upper tidal location and Kenya’s seasonal climate (McClanahan, 1988; Lieth et
al., 1999), the trees that are growing here experience drought periods up to several weeks a month,
associated with rising salinity levels from 34 ‰ (sea water salinity) to 100 ‰ and even higher
during the dry season. In the rainy season, salinity drops to levels approaching fresh water during
the weeks without inundation. At the seaward side of the forest, trees are flooded daily by seawater
of almost constant salinity over the seasons. Trees growing at a different distance from the sea are
thus characterized by an equally different water status, related with different risk levels for the water
transport. A. marina, occurring at both extremes of the coastal slope, reflects the contrasting growth
conditions in its morphology with dwarf trees towards the land and trees of impressive stature towards
the sea. The meaning of water stress for trees is the chance of failure in keeping a continuous water
flow from soil to canopy.
13
General Introduction
In this way intervessel pits at least partly determine the degree of compartmentalization of the water
transport system that is supposed to be balanced against prevailing environmental conditions (Orians
et al., 2004; Ellmore et al., 2006). The mechanism behind drought-induced cavitation is explained by
the air-seeding theory (Sperry and Tyree, 1988; Oertli, 1993; Jarbeau et al., 1995; Salleo et al., 1996)
that is based on the cohesion tension theory by putting a steep tension gradient, caused by soil water
deficit or salinity, at its origin. In the middle of the pit chamber between two vessels, a more or less
porous pit membrane is situated that is composed of remnants of the primary cell walls and the middle
lamella (Fig. 1). According to the theory, an air bubble is seeded when the pressure gradient over the
pit membrane surpasses a critical pressure, determined by the pit membrane pore size and the surface
tension of the gas-water meniscus (Fig. 2a). The air is sucked in from neighbouring already air-filled
vessels that are common in a plant (Tyree et al., 1994). Cycles of cavitation and refilling take place
on a diurnal basis (Cochard et al., 2000; Melcher et al., 2001; Domec et al., 2006b; Woodruff et al.,
2007), in parallel with variations in transpiration rate (Bucci et al., 2003) and also common processes
like leaf abscission and herbivory cause cavitation. Moreover, although spontaneous air-seeding in
xylem sap is extremely rare (Steudle, 2001), it is uncertain at this moment whether gas nucleation
also happens via still other mechanisms than air-seeding at pit membranes and if yes what their
contribution may be in the inactivation of parts of the sapwood (Tyree and Dixon, 1986; Tyree, 1997;
Shen et al., 2002). Freeze-induced cavitation happens by air exsolving from the frozen water when
melting. Intervessel pits are thus not involved in this mechanism (Hacke and Sperry, 2001).
D Air pockets in intervessel pits between a functional and a refilling vessel compartmentalize vessel refilling
(white frame in C). The inclination of the pit chamber walls (2α) and the contact angle between water and ves-
sel wall (θ) determine the strength of the air-water meniscus to disconnect the vessel, under positive pressure,
from the suction force of transpiration until full embolism repair.
14
General Introduction
Embolisms are not necessarily permanent although refilling seems not to be a universal process (Hacke
and Sperry, 2003) and the mechanism is still a matter of debate. The obscurity is in the process of
pressurizing the air-filled vessel while neighbouring vessels are still under tension. In this respect the
importance of vessel associated cells has been mentioned long before (Braun, 1984; Beeckman, 1999).
Osmotic water shifting by secretion of sugars into vessels has even been suggested as a mechanism
for water ascent alternating with transpiration. Although the permanent presence of positive pressures
in the hydrosystem has not found wide support, it agrees with the currently suggested mechanism
for embolism repair. Large osmotically active solutes are supposed to be transported from vessel
associated cells to the embolized vessel. Since the solutes are too large to cross pit membranes water
is pulled in from neighbouring active vessels or parenchyma cells to refill the blocked vessel (Fig. 2b)
(Hacke and Sperry, 2003). In the case the solutes are small (Fig. 2c-d), air-pockets in the intervessel
pits can disconnect the air-filled vessel from the transpiration stream as long as the vessel is not yet
fully refilled (Holbrook and Zwieniecki, 1999). An alternative or perhaps additional mechanism is the
hydrolysis of starch within vessel associated cells leading to pressure imbalances after water uptake
driving water in the embolized vessel (Bucci et al., 2003; Salleo et al., 2004). The water flow could
be directed towards the embolized vessel via activation of specific water channels (De Boer and
Fig. 2 Drought-induced cavitation as explained by the air-seeding theory and mechanisms of embolism repair.
A Air is seeded in from a pore in the pit membrane when a critical pressure difference, between the already
embolized vessel and the functional vessel, is exceeded and the gas-water meniscus breaks. B One proposed
mechanism to refill embolized vessels is based on the transport of large osmotically active solutes from living
vessel associated cells. Because the solutes are too big to cross pit membranes water is attracted from both
the transpiration stream and neighbouring cells. The inflowing water creates a positive pressure in the refilling
vessel, forcing the air back into solution (Fig. adapted from Hacke and Sperry, 2003). C An alternative mecha-
nism of embolism repair depends on small solutes that also draw in water from neighbouring vessels but not
from adjacent functional vessels. Since these solutes can cross pit membranes the refilling vessel should be
disconnected from the transpiration stream to create a positive pressure. (continued on p. 14)
15
General Introduction
Volkov, 2003). The trigger for starch hydrolysis might be vibration of the vessel walls upon cavitation
(Salleo et al., 2004; Salleo, 2006). Although at present there is no conclusive evidence for one or a
combination of the presented strategies (Stiller et al., 2005), all mechanisms are conclusive about
the metabolic control over the process of embolism repair. Living tissue such as parenchyma and
phloem (Salleo et al., 1996; Salleo et al., 2004; Salleo, 2006) seem to be the basic requirement for
refilling to take place, which can happen in a time range of minutes (Facette et al., 2001; Zwieniecki
et al., 2001a; Salleo et al., 2004). Under natural drought stress conditions, however, it could take days
(Hacke and Sperry, 2003).
Cycles of cavitation and repair can not go on infinitely since the refilling process slows down (Lee
and Kim, 2007) and since cavitation fatigue takes place reducing cavitation resistance (Hacke et al.,
2001b). This weakening is most likely caused by irreversible membrane damage after stretching
during cavitation (Domec et al., 2006a). It could be minimized by fibers acting as buttresses (Jacobsen
et al., 2005) although this could not be supported in a more recent study (Pratt et al., 2007). It is yet
unknown whether it concerns a permanent state but in each case there is a moment embolism repair
does no longer take place and vessels become permanently blocked where after they can be filled
with gums, tyloses (Hargrave et al., 1994) or mucilages (Ewers et al., 2004). More and more studies,
however, prove cavitation to be part of a hydraulic signal to regulate the water status of the tree
rather than being an unavoidable evil, associated with the transport of water under tension. Cavitated
vessels could function as water storage tissue (Stratton et al., 2000; Melcher et al., 2001). Older xylem
vessels with a lower cavitation resistance might secure optimal functioning of the younger sapwood
by acting as a water reservoir (Melcher et al., 2003). Spatial partitioning of xylem vulnerability to
the most distal parts of the tree can have the advantage of sacrificing less important or more easily
refilled parts of the tree, saving and mitigating tension in surviving more basally oriented branches
(Hacke and Sperry, 2001; Choat et al., 2005b; Domec et al., 2006b; Woodruff et al., 2007). It follows
that the hydraulic system is not simply generated to avoid gas nucleation but to reach the most
beneficial activity state. Therefore, trees adapt both structurally and physiologically to the prevailing
environmental conditions.
Approach: the added value of ecological wood anatomy in clarifying tree functioning
The adaptations of trees to the environment, with its spatial and temporal variations, are situated on
three related levels: tree morphology, tree physiology and wood anatomy (Maseda and Fernández,
2006). To understand the findings of physiological studies a profound understanding of the hydraulic
basis, the structure of the water transport system, is required (e.g. Meinzer, 2003; Choat et al., 2005a;
Domec et al., 2007; Loepfe et al., 2007). The direct relationship between hydraulic structure and
water transport is reflected in its adaptations to changing stress conditions within and between trees
and species growing under different environmental conditions.
In the formation of their water transport system trees strive to an optimal compromise between
efficiency and safety (Wheeler et al., 2005; Hacke et al., 2006; Sperry et al., 2008). The trade-
off results from the fact that a high efficiency to transport water brings along a facilitated spread
and expansion of gas bubbles, blocking the water flow in sometimes extended parts of the conduit
network. Across distantly related species no or only a weak trade-off was observed (Maherali et al.,
2004), which might be explained by ample strategies to balance the water status of the tree, next to
structural adaptations of the hydrosystem (Loepfe et al., 2007). Tree morphology and tree physiology
come also into play when extending to the whole-tree level (e.g. Melcher et al., 2001; Cruiziat et
al., 2002; Kocacinar and Sage, 2004; Bréda et al., 2006; Junghans et al., 2006; Krauss et al., 2006b;
Sobrado and Ewe, 2006). In addition, one should keep in mind that the hydraulic architecture is also
16
General Introduction
determined by internal factors such as the genetic background of the tree (Baas et al., 1983; February
and Manders, 1999; Reich et al., 2003; Dünisch et al., 2004) and its mechanical requirements (Taneda
and Tateno, 2004) although there should not necessarily be a trade-off with transport needs (Woodrum
et al., 2003; Jacobsen et al., 2005; Sperry et al., 2006).
Here, we will discuss how the hydraulic structure can be adjusted to meet the safety requirements for
sap flow under water stress. Since no freezing occurs at the Kenyan coast, only adaptations to drought
stress will be discussed and this from the tissue level, over the cellular level to the sub-cellular level
of the intervessel pits.
17
General Introduction
Fig. 3 Secondary growth via successive cambia in Avicennia marina. A Transverse section of a portion of
the outermost stem wood. Three cambia are simultaneously active (arrow heads), forming xylem (X) to the
inside, sclereids (SC) and phloem (P) to the outside. At the barkside a new cambium has just developed. More
inward, part of the xylem has not yet lignified. In the innermost growth layer, the parenchyma tissue (PA) in
between the differentiating phloem strands (Ps) started lignification. Scale bar, 100 μm. B Transverse section
of a seedling where the alternating xylem and phloem bands are clearly visible. The phloem bands consist of
phloem strands embedded in a layer of parenchyma and delineated from the next growth layer by two to three
cell layers of sclereids. Scale bar, 1000 μm. C Sanded stem disc with the phloem and xylem bands appearing
as light and dark coloured bands, respectively. Scale bar, 1000 μm. D Detail of the outermost stem wood indi-
cated by the frame in A. A new cambium has recently developed in the parenchyma layer to the outside of the
sclereids. Scale bar, 100 μm. Double staining with safranin-fast green.
small-scale studies underscore their findings and characterize a relatively safe hydraulic structure by
vessels of a small diameter (Hargrave et al., 1994; Jarbeau et al., 1995; Villar-Salvador et al., 1997;
Corcuera et al., 2004; Junghans et al., 2006; Sobrado, 2007), composed of short vessel elements (Sun
and Lin, 1997; Villar-Salvador et al., 1997), joined together into short vessels (Sperry et al., 1988),
occurring in high frequency (Choat et al., 2005a; Verheyden et al., 2005; Sobrado, 2007), in vessel
groups rather than solitary (López et al., 2005) and with a thick vessel wall (Sperry et al., 1988).
The link between vessel diameter and cavitation resistance can be explained in two ways by the
indirect influence, clarifying the flexibility of the correlation (Tyree et al., 1994; Mauseth and Plemons-
Rodriguez, 1998; Hacke and Sperry, 2001; Sperry et al., 2006), of vessel size on the air-seeding
process. First, small vessels have an equally small wall area, resulting in a relatively low pit area per
vessel and thus cavitation risk (Choat et al., 2005b; Hacke et al., 2006). The correlation between the
total pit membrane area and the cavitation vulnerability could be explained by the increased chance
18
General Introduction
on an exceptionally large pit membrane pore, prone to air-seeding (Hargrave et al., 1994; Wheeler
et al., 2005; Hacke et al., 2006). Second, vessels with a small diameter have a large surface-to-
volume ratio which would aid in controlling the xylem sap composition (De Boer and Volkov, 2003).
Given the ion regulation of the hydraulic conductance via hydrogels in pit membrane pores (see
section ‘Approach’, ‘The sub-cellular level’) this could be of adaptive value under stress conditions.
Recently, another positive effect of ions on sap flow was assigned to the prevention of coalescence
rather than the development of air bubbles (Canny et al., 2007). The same authors presented a new
theory saying that perforation plates of small vessels are more likely to trap cell wall residues from
vessel element endings after hydrolysis that can act as gass nucleation sites. In this way small vessels
could trap exsolved air, leaving degassed less cavitation-prone water for more efficient flow through
wide vessels. This theory would only apply if cavitation triggered by constrictions along the flow
path, occurs more easily than air seeding to which large vessels are more vulnerable. But in the first
place, future research is challenged to underscore the formation of air bubbles via irregularities of
the vessel wall next to air-seeding. This would have serious implications for the current views on
structure-function relationships. Species with scalariform perforation plates, like Rhizophora, would
inherently be more vulnerable to cavitation and we should have to revise our vision on the functional
advantage of a bimodal distribution in vessel size, so often observed in arid environments (Baas et
al., 1983).
In addition to reducing the air-seeding probability, small vessels mitigate the negative effect of it.
Simply because of their small size, they maximize the proportion of active xylem upon hydraulic
failure. The effect is enhanced by the correlation between vessel diameter and vessel length, across
species (Sperry et al., 2005; Hacke et al., 2006) and within ring porous species when comparing
latewood and earlywood (Zimmermann and Jeje, 1981). Reducing vessel element length (Baas et
al., 1983), and especially total vessel length (Comstock and Sperry, 2000; Hacke and Sperry, 2001)
hinders the spreading of embolisms by perforation plates or vessel endings, respectively. The parts
of the conducting network that are blocked can be readily circumvented if vessel density and vessel
grouping are high, providing more alternative routes to circumvent air bubbles (Villagra and Roig
Juñent, 1997; Cruiziat et al., 2002; Kitin et al., 2004). Besides, the strategy of packing as many
vessels as possible in the xylem guarantees a basic hydraulic conductance even after considerable
losses to embolism (Hacke et al., 2006). The higher redundancy of the vessel network also implies a
higher potential for air bubbles to spread to other vessels highlighting the complexity of studying the
functional significance of the hydraulic structure (Loepfe et al., 2007). With respect to the connectivity
dilemma, geometry of intervessel pits and wall chemistry should not be overlooked (Fig. 2d). The
pressure needed to break the gas-water interface in the pit chamber between a functional and an
embolized vessel depends on the contact angle between water and vessel wall, and the inclination
of the pit chamber walls (Holbrook and Zwieniecki, 1999; Zwieniecki and Holbrook, 2000; Meyra
et al., 2007). Gas crosses pit membranes thus less readily than water, making it easier to understand
the advantages of a more integrated transport system. It provides not only the opportunity to bypass
embolisms but also to exploit spatially localized soil resources (Orians et al., 2004; Ellmore et al.,
2006), outweighing the potential disadvantage of an intense packing of vessels.
The increased xylem tension does not only lead to deactivation of vessels by embolization but also
by vessel implosion. This can be hindered by more sturdy fibers to support vessels (Jacobsen et
al., 2005; Cochard et al., 2007; Jacobsen et al., 2007b) or directly by increasing the ratio of the
intervessel wall thickness to the vessel lumen diameter (Hacke et al., 2001a) that is related to wood
density (Preston et al., 2006; Jacobsen et al., 2007a). It should be noted that between more related
species the relationship between wood density and vulnerability to conductive failure seems to be
missing (Cochard et al., 2007). Besides, the fact that a higher wood density results in a lower water
storage capacity (Domec et al., 2006b) emphasizes the importance of integrative studies, looking at
19
General objective & hypotheses
the hydraulic architecture as a whole since it is the balance between all characteristics that determines
the cavitation vulnerability of the tree.
The relationship between pit membrane structure and cavitation resistance shows some flexibility. A
first explanation is given by the cavitation resistance that is determined by the maximum and not the
average pore diameter of the pit membrane. If air seeding would take place at a random pore, pit
membranes with a small mean pore size would have an increased cavitation resistance (Zimmermann,
1983; Sperry and Tyree, 1988). However, if air seeding would first take place at the largest pore
per vessel, average pit membrane area per vessel would be the key factor determining cavitation
resistance. The larger the pit area, the bigger the chance on an exceptional large pore increasing
cavitation vulnerability (Wheeler et al., 2005; Hacke et al., 2006; Choat et al., 2008). This so-called
pit area hypothesis is the most supported option at the moment. The size of the maximum pore is
unrelated to the average pore diameter and is the result of pit membrane damage or stretching (Choat
et al., 2003). The difficulty in observing these large, first air-seeding pores might be related to the
temporary condition of their exceptional size (Choat et al., 2004). Alternatively, the large pore is in
its natural state but rare (Wheeler et al., 2005). Next to the low frequency of these exceptionally large
pores, their identification as the pores causing cavitation is hampered by the high risk for artefacts
inherent to the delicate pit membranes (Choat et al., 2003; Choat et al., 2006). The stretching or even
rupture of the pit membrane when aspirated by the pressure difference between a functional and an
embolized vessel (Hacke et al., 2001b; Choat et al., 2003) gives a second clarification for the flexible
correlation between pit membrane porosity and cavitation resistance. Several strategies have evolved
to prevent deviation of the porosity from its pristine state. The variation observed between species
in pit membrane thickness (Schmid and Machado, 1968; Sano, 2005; Choat et al., 2008) could
translate in varying probability to stretch excessively and develop large pores. Next to changes of
the pit membrane itself, vestures or appendices of the vessel wall in pit canals and sometimes also
pit chambers (Fig. 4a), impede excessive stretching by supporting the pit membrane (Fig. 4b) and
hence lowering the vulnerability for drought induced embolism (Zweypfenning, 1978; Jansen et al.,
1998; Jansen et al., 2003; Choat et al., 2004; Jansen et al., 2004; Sperry and Hacke, 2004). Thirdly,
next to structural adjustments, cavitation resistance can be affected by pit membrane chemistry
influencing the air-water interface (see section ‘Approach’, ‘The cellular level’). Besides, pore size
can be regulated by changes in the chemical composition of the xylem sap although its significance
in planta remains to be demonstrated. Ion-regulation of pit membrane pore size is based on the
hydrogel nature of pectins in the pit membrane (Tibbits et al., 1998; Ridley et al., 2001). Depending
on the ionic composition of the xylem fluid, pectins swell under low ion content and shrink under high
ion content when cross-linkages are being built. Adjusting hydraulic conductivity by changing ionic
content provides an opportunity to compensate for the negative effect of embolisms (Zwieniecki et
al., 2001b; Lopez-Portillo et al., 2005; van Ieperen, 2007). In this respect, less lignified vessels would
offer the highest benefit. Because of the more intimate contact between pectins and ions, the hydraulic
20
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työskenneltiin. Mutta ihmeellistä on, että tämä hallitus, joka
sisäoloissa oli täynnä intohimoisia puolueriitoja, ja jonka päivästä
päivään täytyi keksiä uusia keinoja vastustaakseen kunnialla
ulkovihollista, tahtoi ja voi uhrata niin paljon kallista aikaansa ja
kalliita voimiansa hankkiakseen Ranskalle uusia kultuurivoittoja. Että
se todella niin teki, todistaa enemmän kuin mikään muu, että tämä
hallitus ei ollut mikään ilkimysten ja pyövelien kokous, kuten usein
tahdotaan väittää, vaan että sillä oli käytettävänään paljon
valistusta, paljon hyvää tahtoa, suunnaton työkyky ja suuri rakkaus
rauhallisiin harrastuksiin ja rauhaisaan työhön. Ei halu käytännössä
toteuttaa jotakin oppia tai järjestelmää, vaan pääasiallisesti
epätoivoiset sisälliset ja ulkonaiset olosuhteet ne johtivat konventin
väkivaltaisuuksiin, kiihkoiluun ja sortoon.
*****
*****
Jos luomme silmäyksen jälellepäin edellä kosketeltuihin konventin
reformeihin, näkyy selvästi että konventti useimmassa niistä piti
silmällä pedagogista päämäärää. Kasvatus oli yksi vallankumouksen
iskusanoja, se oli myös vallankumouksen etevimpiä päämaaleja.
Kasvatus — eikä ainoasti vapauteen, tasa-arvoisuuteen,
kansalaisuuteen ja tasavaltaisuuteen, vaan myös oppiin ja tietoihin,
vakaasen työhön ja tosi sivistykseen. Ei ollut siis ihmettä, että
sellaisissa olosuhteissa kouluopetuksen järjestäminen oli
vallankumouksen rakkaimpia toimia ja kenties se kysymys, joka
kauimmin ja sitkeimmin antoi vallankumousmiehille ajattelun aihetta.
Kaksi syytä oli lisäksi jo alusta pitäen auttanut katseiden
suuntautumista tälle alalle — ensiksi se tavattoman kehno tila, jossa
kouluopetus oli vallankumouksen puhjetessa, toiseksi ne mahtavat
herätteet, joita valistusfilosofit olivat antaneet kasvatuksen ja
opetuksen täydelliseksi uudestimuodostamiseksi. Rousseau oli
kirjallaan Emile ou l'éducation rikkonut välit kaikkien vanhojen
traditsionien ja kaikkien totuttujen tapojen kanssa ja esittänyt
luonnollisen kehityksen ja ihmisellisen hyvyyden periaatteihin
nojautuvan ihanteen. Sellainen oli myös se ihanne, jonka
vallankumous jo alusta asti asetti itselleen.
*****
"Homme libre et pensant, né pour haïr les rois, N'aimer que mes
égaux et servir ma patrie, Vivre de mon travail ou de mon industrie,
Abhorrer l'esclavage et me soumettre aux lois." ["Vapaa ja ajatteleva
ihminen, syntynyt vihaamaan kuninkaita, rakastamaan ainoastaan
vertaisiani ja palvelemaan isänmaatani, elämään työstäni ja
toimeliaisuudestani, kammoamaan orjuutta ja alistumaan lakien
alaiseksi."]
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