Choanoflagellate and choanocyte collar-flagellar systems and the assumption of homology.

The similarities between the choanoflagellates and the choanocytes of sponges have been discussed for more than a century yet few studies allow a direct comparison of the two. We reviewed current knowledge of the collar and flagellum and compared their structure and function in the choanoflagellate Monosiga brevicollis and the sponge Spongilla lacustris. Collar microvilli were of similar length and number, but the shape of the collar differed between the two cells. In Monosiga, collars were flared and microvilli were joined by a single band of glycocalyx mid-way along their length; in Spongilla, collars formed a tube and microvilli were joined by a mesh of glycocalyx. Monosiga flagella beat at least four times faster than those in Spongilla. Flagellar vanes were found in both cell types. In both cells, the flagella and so probably also the vanes maintained moving points of contact with the microvilli, which suggested that collars and flagella were integrated systems rather than independent units. There were fundamental differences in how the collar and flagella interacted, however. In Spongilla, the flagellum bent upon contact with the collar; the flagellar amplitude was fitted to the collar diameter. In Monosiga, the flagellar amplitude was unaffected by the collar; instead the collar diameter appeared fitted to the flagellum. These differences suggest that though choanocytes and choanoflagellates are similar, homology cannot be taken for granted. Similarities in collar-flagellum systems separated by 600 million years of evolution, whether maintained or convergent, suggest that these form important adaptations for optimizing fluid flow through micro-scale filters.

Frost fatigue and spring recovery of xylem vessels in three diffuse-porous trees in situ.

Frost has been shown to cause frost fatigue (reduced cavitation resistance) in branch segments in the lab. Here, we studied the change in cavitation resistance and percent loss of conductivity (PLC) from fall to spring over 2 consecutive years in three diffuse-porous species in situ. We used the cavitron technique to measure P25 , P50 and P90 (the xylem pressure causing a 25, 50 and 90% conductivity loss) and PLC and stained functioning vessels. Cavitation resistance was reduced by 64-87% (in terms of P50 ), depending on the species and year. P25 was impacted the most and P90 the least, changing the vulnerability curves from s- to r-shaped over the winter in all three species. The branches suffered an almost complete loss of conductivity, but frost fatigue did not necessarily occur concurrently with increases in PLC. In two species, there was a trade-off between conduit size and vulnerability. Spring recovery occurred by growth of new vessels, and in two species by partial refilling of embolized conduits. Although newly grown and functioning conduits appeared more vulnerable to cavitation than year-old vessels, cavitation resistance generally improved in spring, suggesting other mechanisms for partial frost fatigue repair.

Does freezing and dynamic flexing of frozen branches impact the cavitation resistance of Malus domestica and the Populus clone Walker?

Frost damage to the xylem conduits of trees is a phenomenon of eco-physiological importance. It is often documented in terms of the percentage loss of conductivity (PLC), an indicator of air filling of the conduits. However, trees that refill their conduits in spring could be impacted more by damage to the conduits that reduce cavitation resistance, making them more susceptible to future drought events. We investigated whether ice formation, dynamic flexing of frozen branches or freeze-thaw events could reduce the cavitation resistance (cause "frost fatigue") in first-year shoots of apple (Malus domestica) and clonal hybrid cottonwood (Walker). Frost fatigue was measured in terms of P50 (the negative xylem pressure required to cause a 50 % loss of conductivity). All treatment groups showed significant frost fatigue, with the exception of the pre-flushed, constantly frozen poplar branches. The P50 following freeze treatments was approximately 50 % of the pre-freeze values. The effect tended to be greater in freeze-thawed branches. Dynamic bending of the branches had no effect on either PLC or P50. In three out of four cases, there was a significant correlation between P50 and PLC. Frost fatigue occurred in both apple and poplar, two unrelated species with different drought and frost tolerances, suggesting that it may be a widespread phenomenon that could impact the ecophysiology of temperate forests.

Changes in hydraulic conductivity, mechanical properties, and density reflecting the fall in strain along the lateral roots of two species of tropical trees.

Roots have been described as having larger vessels and so greater hydraulic efficiency than the stem. Differences in the strength and stiffness of the tissue within the root system itself are thought to be an adaptation to the loading conditions experienced by the roots and to be related to differences in density. It is not known how potential mechanical adaptations may affect the hydraulic properties of the roots. The change in strength, stiffness, conductivity, density, sapwood area, and second moment of area distally along the lateral roots of two tropical tree species in which the strain is known to decrease rapidly was studied and the values were compared with those of the trunk. It was found that as the strain fell distally along the roots, so did the strength and stiffness of the tissue, whereas the conductivity increased exponentially. These changes appeared to be related to differences in density. In contrast to the distal-most roots, the tissue of the proximal roots had a lower conductivity and higher strength than that of the trunk. This suggests that mechanical requirements on the structure rather than the water potential gradient from roots to branches are responsible for the general pattern that roots have larger vessels than the stem. In spite of their increased transectional area, the buttressed proximal roots were subjected to higher levels of stress and had a lower total conductivity than the rest of the root system.

Complex flagellar motions and swimming patterns of the flagellates Paraphysomonas vestita and Pteridomonas danica.

Most flagellates with hispid flagella, that is, flagella with rigid filamentous hairs (mastigonemes), swim in the direction of the flagellar wave propagation with an anterior position of the flagellum. Previous analysis was based on planar wave propagation showing that the mastigonemes pull fluid along the flagellar axis. In the present study, we investigate the flagellar motions and swimming patterns for two flagellates with hispid flagella: Paraphysomonas vestita and Pteridomonas danica. Studies were carried out using normal and high-speed video recording, and particles were added to visualize flow around cells generating feeding currents. When swimming or generating flow, P. vestita was able to pull fluid normal to, and not just along, the flagellum, implying the use of the mastigonemes in an as yet un-described way. When the flagellum made contact with food particles, it changed the flagellar waveform so that the particle was fanned towards the ingestion area, suggesting mechano-sensitivity of the mastigonemes. Pteridomonas danica was capable of more complex swimming than previously described for flagellated protists. This was associated with control of the flagellar beat as well as an ability to bend the plane of the flagellar waveform.

Surface tension phenomena in the xylem sap of three diffuse porous temperate tree species.

In plant physiology models involving bubble nucleation, expansion or elimination, it is typically assumed that the surface tension of xylem sap is equal to that of pure water, though this has never been tested. In this study we collected xylem sap from branches of the tree species Populus tremuloides, Betula papyrifera and Sorbus aucuparia over 3 months. We measured the instantaneous surface tension and followed changes over a period of 0.5-5 h using the pendant drop technique. In all three species the instantaneous surface tension was equal to or within a few percent of that of pure water. Further, in B. papyrifera and S. aucuparia the change over time following drop establishment, although significant, was very small. In P. tremuloides, however, there was a steep decline in surface tension over time that leveled off towards values 21-27% lower than that of pure water. This indicated the presence of surfactants. The values were lower for thinner distal branch segments than for proximal ones closer to the trunk. In some species it appears valid to assume that the surface tension of xylem sap is equal to that of water. However, in branch segments of P. tremuloides close to the terminal bud and hence potentially in other species as well, it may be necessary to take into account the presence of surfactants that reduce the surface tension over time.

Interrelations between hydraulic and mechanical stress adaptations in woody plants.

The fields of plant water relations and plant biomechanics have traditionally been studied separately even though often the same tissues are responsible for water transport and mechanical support. There is now increasing evidence that hydraulic and mechanical adaptations may influence one another. We studied the changes in the hydraulic and mechanical properties of the wood along lateral roots of two species of buttressed trees. In these roots, the mechanical contstraints quantified by strain measurements are known to decrease distally. Further, we investigated the effect of mechanical loading on the vessel anatomy in these and four other species of tropical trees. We found that as the strain decreased, the wood became progressively less stiff and strong but the conductivity increased exponentially. This was reflected in that adaptations towards re-enforcing mechanically loaded areas resulted in xylem with fewer and smaller vessels. In addition a controlled growth experiment on three tree species showed that drought adaptation may results in plants with stronger and stiffer tissue. Our results indicate that hydraulic and mechanical stress adaptations may be interrelated, and so support recent studied suggesting that physiological responses are complex balances rather than pure optimisations.

Changes in vessel anatomy in response to mechanical loading in six species of tropical trees.

It is well known that trees adapt their supportive tissues to changes in loading conditions, yet little is known about how the vascular anatomy is modified in this process. We investigated this by comparing more and less mechanically loaded sections in six species of tropical trees with two different rooting morphologies. We measured the strain, vessel size, frequency and area fraction and from this calculated the specific conductivity, then measured the conductivity, modulus of elasticity and yield stress. The smallest vessels and the lowest vessel frequency were found in the parts of the trees subjected to the greatest stresses or strains. The specific conductivity varied up to two orders of magnitude between mechanically loaded and mechanically unimportant parts of the root system. A trade-off between conductivity and stiffness or strength was revealed, which suggests that anatomical alterations occur in response to mechanical strain. By contrast, between-tree comparisons showed that average anatomical features for the whole tree seemed more closely related to their ecological strategy.