Alternative fire resistance strategies in savanna trees

Bark properties (mainly thickness) are usually presented as the main explanation for tree survival in intense fires. Savanna fires are mild, frequent, and supposed to affect tree recruitment rather than adult survival: trunk profile and growth rate of young trees between two successive fires can also affect survival. These factors and fire severity were measured on a sample of 20 trees near the recruitment stage of two savanna species chosen for their contrasted fire resistance strategies (Crossopteryx febrifuga and Piliostigma thonningii). Crossopteryx has a higher intrinsic resistance to fire (bark properties) than Piliostigma: a 20-mm-diameter stem of Crossopteryx survives exposure to 650°C, while Piliostigma needs a diameter of at least 40 mm to survive. Crossopteryx has a thicker trunk than Piliostigma: for two trees of the same height, the basal diameter of Crossopteryx will be 1.6 times greater. Piliostigma grows 2.26 times faster than Crossopteryx between two successive fires. The two species have different fire resistance strategies: one relies on resistance of aboveground structures to fire, while the other relies on its ability to quickly re-build aboveground structures. Crossopteryx is able to recruit in almost any fire conditions while Piliostigma needs locally or temporarily milder fire conditions. In savannas, fire resistance is a complex property which cannot be assessed simply by measuring only one of its components, such as bark thickness. Bark properties, trunk profile and growth rate define strategies of fire resistance. Fire resistance may interact with competition: we suggest that differences in fire resistance strategies have important effects on the structure and dynamics of savanna ecosystems. Received: 16 August 1996 / Accepted: 4 January 1997     

Drip irrigation,soil characteristics and the root distribution and root activity of olive trees

A study was carried out on the root distribution and root activity of the olive tree (Olea Europaea, L., var. manzanillo) as influenced by drip irrigation and by several soil characteristics such as texture and depth. The experiments were conducted in two plots within a drip-irrigated grove of 20-year-old trees planted at 7×7 m spacing. One soil was a sandy loam, the other a clay-loam. Both cylinder and trench methods were used to determine root distribution. Labelling with ³²P was used to determine root activity. Under dryland conditions the adult tree adapted its rooting system, following the installation of a drip system, by concentrating the roots within the wet soil zones near the drippers. The highest root densities occur in those zones, down to a 0.6 m depth, the most abundant being the <0.5 mm diameter roots. The most intensive root activity was also found in that zone. For a given irrigation system, wet soil bulbs are more extensive and therefore root distribution expands to a larger soil volume when the soil is more clayey and with a hard calcareous pan present at about 0.8 m depth which prevents deep drainage.     

Wind breakage of trees initiated by root delamination

Summary There are different types of tree failure under wind loading. Here a special mechanism is discussed by which redirectioning of the tensile forces acting in the tree stem causes a lateral force to become effective away from the stem-root transition contour line. This force leads to delamination of the root followed by development of an axial crack running up the stem, until total stem failure by fibre buckling on the compression side of the stem completes the destruction. It is shown that buttress roots could reduce the danger associated with this failure mechanism.     

Transpiration and root water uptake by olive trees

While the cultivated olive tree (Olea europaea L.) is known to be sclerophyllous and effective at tolerating drought, little is known of its short-term water-use dynamics for most studies have been based on longer-term, water-balance information. We present here, for the first time, heat-pulse measurements of the sap flux measured not only within the semi-trunk of an olive tree, but also within a root excavated close to the stump. One tree in the olive grove near Seville in Spain had regularly received basin irrigation during the summer, whereas the other, growing on this deep silt loam, had been without water for over 3 months. Following a flood irrigation of 730 L to a dyked area around the tree, the regularly-irrigated olive maintained a transpiration rate of 1.65 mm³ mm^–2 d^–1, on a leaf area basis, for only 3 days following the irrigation. This rate was maintained for a total consumption of 110 L. It then began again to limit its rate of water use with transpiration falling below that predicted for well-watered conditions by the Penman-Monteith equation. The flow of sap in the near-surface root dropped concomitantly. Meanwhile the unirrigated tree was using water at just 0.78 mm d^–1. Yet following an irrigation of 870 L it only lifted its consumption to 1.12 mm d^–1, on a leaf area basis. Neither did it recover its leaf water potential following this wetting because of an inability to refill cavitated vessels. These data again show olive to be a parsimonious and cautious consumer of soil water.     

Long range allosteric control of cytoplasmic dynein ATPase activity by the stalk and C-terminal domains

The dynein motor domain consists of a ring of six AAA domains with a protruding microtubule-binding stalk and a C-terminal domain of unknown function. To understand how conformational information is communicated within this complex structure, we produced a series of recombinant and proteolytic rat motor domain fragments, which we analyzed enzymatically. A recombinant 210-kDa half-motor domain fragment surprisingly exhibited a 6-fold higher steady state ATPase activity than a 380-kDa complete motor domain fragment. The increased ATPase activity was associated with a complete loss of sensitivity to inhibition by vanadate and an approximately 100-fold increase in the rate of ADP release. The time course of product release was discovered to be biphasic, and each phase was stimulated approximately 1000-fold by microtubule binding to the 380-kDa motor domain. Both the half-motor and full motor domain fragments were remarkably resistant to tryptic proteolysis, exhibiting either two or three major cleavage sites. Cleavage near the C terminus of the 380-kDa motor domain released a 32-kDa fragment and abolished sensitivity to vanadate. Cleavage at this site was insensitive to ATP or 5'-adenylyl-beta,gamma-imidodiphosphate but was blocked by ADP-AlF3 or ADP-vanadate. Based on these data, we proposed a model for long range allosteric control of product release at AAA1 and AAA3 through the microtubule-binding stalk and the C-terminal domain, the latter of which may interact with AAA1 to close the motor domain ring in a cross-bridge cycle-dependent manner.     

Increasing temperatures can improve seedling establishment in arid-adapted savanna trees

Plant species are shifting their ranges in response to global climate change, thus intensifying the need to predict such changes accurately. As the environmental requirements controlling plant distribution act differently at each developmental stage, there is a need to acquire a demographic-specific understanding of the factors which determine these distributions. Here we investigated the germination niche of two common savanna species Acacia nigrescens and Colophospermum mopane, with the aims to disentangle the direct and indirect effects of temperature on seed germination and establishment and to explore the impact of higher temperatures on the establishment success of savanna trees. Under laboratory conditions, we used thermal gradient plates to determine the thermal germination niche of both species, and a water stress experiment was conducted on C. mopane to account for water–temperature interactions. Using these data we parameterised a soil-moisture model to determine germination and establishment success under field conditions at current and future temperatures (+4 °C). Based on this model, higher future temperatures will not limit germination directly, but they will reduce the number of germination events by reducing the time window of suitable available soil water. Conversely, warmer conditions will accelerate the rate of radicle extension and increase the frequency of seedling establishment events. An additional advantage of higher temperatures is that fewer seeds will germinate, resulting in slower seed bank depletion when successful seedling establishment events do occur.     

Long range molecular dynamics study of regulation of eukaryotic glucosamine-6-phosphate synthase activity by UDP-GlcNAc

Glucosamine-6-phosphate (GlcN-6-P) synthase catalyses the first and practically irreversible step in hexosamine metabolism. The final product of this pathway, uridine 5’ diphospho N-acetyl-D-glucosamine (UDP-GlcNAc), is an essential substrate for assembly of bacterial and fungal cell walls. Moreover, the enzyme is involved in phenomenon of hexosamine induced insulin resistance in type II diabetes, which makes it a potential target for antifungal, antibacterial and antidiabetic therapy. The crystal structure of the isomerase domain of GlcN-6-P synthase from human pathogenic fungus Candida albicans, in complex with UDP-GlcNAc has been solved recently but it has not revealed the molecular mechanism of inhibition taking place under UDP-GlcNAc influence, the unique feature of the eukaryotic enzyme. UDP-GlcNAc is a physiological inhibitor of GlcN-6-P synthase, binding about 1 nm away from the active site of the enzyme. In the present work, comparative molecular dynamics simulations of the free and UDP-GlcNAc-bounded structures of GlcN-6-P synthase have been performed. The aim was to complete static X-ray structural data and detect possible changes in the dynamics of the two structures. Results of the simulation studies demonstrated higher mobility of the free structure when compared to the liganded one. Several amino acid residues were identified, flexibility of which is strongly affected upon UDP-GlcNAc binding. Importantly, the most fixed residues are those related to the inhibitor binding process and to the catalytic reaction. The obtained results constitute an important step toward understanding of mechanism of GlcN-6-P synthase inhibition by UDP-GlcNAc molecule.     

Cadmium stress suppresses lateral root formation by interfering with the root clock

A biological clock activated by oscillating signals, known as root clock, has been linked to lateral root (LR) formation and is essential for regular LR spacing along the primary root. However, it remains unclear how this internal mechanism is influenced by environmental factors known to affect the LR pattern. Here, we report that excessive cadmium (Cd) inhibits LR formation by disrupting the lateral root cap (LRC)‐programmed cell death (PCD)‐regulated root clock. Cd restricts the frequency of the oscillating signal rather than its amplitude. This could be attributed to the inhibition on meristematic activity by Cd, which resulted in decreased LRC cell number and LRC‐PCD frequency. Genetic evidence further showed that LRC cell number is positively correlated with root resistance to Cd. Our study reveals root cap dynamics as a novel mechanism mediating root responses to Cd, providing insight into the signalling pathways of the root clock responding to environmental cues.     

Attack frequency and the tolerance to herbivory of Neotropical savanna trees

Tolerance is the ability of a plant to regrow or reproduce following damage. While experimental studies typically measure tolerance in response to the intensity of herbivory (i.e., the amount of leaf tissue removed in one attack), the impact of how many times plants are attacked during a growing season (i.e., the frequency of damage) is virtually unexplored. Using experimental defoliations that mimicked patterns of attack by leaf-cutter ants (Atta spp.), we examined how the frequency of herbivory influenced plant tolerance traits in six tree species in Brazil’s Cerrado. For 2 years we quantified how monthly and quarterly damage influenced individual survivorship, relative growth rate, plant architecture, flowering, and foliar chemistry. We found that the content of leaf nitrogen (N) increased among clipped individuals of most species, suggesting that Atta influences the allocation of resources in damaged plants. Furthermore, our clipping treatments affected tree architecture in ways thought to promote tolerance. However, none of our focal species exhibited a compensatory increase in growth (increment in trunk diameter) in response to herbivory as relative growth rates were significantly lower in clipped than in unclipped individuals. In addition, the probability of survival was much lower for clipped plants, and lower for plants clipped monthly than those clipped quarterly. For plants that did survive, simulated herbivory dramatically reduced the probability of flowering. Our results were similar across a phylogenetically distinct suite of species, suggesting a potential extendability of these findings to other plant species in this system.     

Simply the best: the transition of savanna saplings to trees

Tree cover in savannas is determined as much by disturbances from fire and herbivory as by rainfall and soil resources. Fire especially acts to limit tree cover via a demographic bottleneck, limiting the recruitment of tree saplings to adults. Because sapling growth rates determine rates of sapling to tree recruitment, predicting changes in tree cover requires data on sapling growth rates, commonly expressed as population means. Here, we discuss the variability in sapling growth rates in Acacia populations in a savanna in Hluhluwe iMfolozi Park in South Africa. Saplings growing at mean rates under typical fire regimes in African savannas would likely never escape the fire‐trap to become adults. Only the fastest growing saplings could grow above the flame zone between fires. We suggest that maximum growth rates are more ecologically relevant than mean growth rates in natural populations and experiments. Maximum growth rates are better than mean growth rates as predictors of sapling release within species, as shown here, and probably of which species are likely ‘winners’ in savanna tree communities.     

Impacts of savanna trees on forage quality for a large African herbivore

Recently, cover of large trees in African savannas has rapidly declined due to elephant pressure, frequent fires and charcoal production. The reduction in large trees could have consequences for large herbivores through a change in forage quality. In Tarangire National Park, in Northern Tanzania, we studied the impact of large savanna trees on forage quality for wildebeest by collecting samples of dominant grass species in open grassland and under and around large Acacia tortilis trees. Grasses growing under trees had a much higher forage quality than grasses from the open field indicated by a more favourable leaf/stem ratio and higher protein and lower fibre concentrations. Analysing the grass leaf data with a linear programming model indicated that large savanna trees could be essential for the survival of wildebeest, the dominant herbivore in Tarangire. Due to the high fibre content and low nutrient and protein concentrations of grasses from the open field, maximum fibre intake is reached before nutrient requirements are satisfied. All requirements can only be satisfied by combining forage from open grassland with either forage from under or around tree canopies. Forage quality was also higher around dead trees than in the open field. So forage quality does not reduce immediately after trees die which explains why negative effects of reduced tree numbers probably go initially unnoticed. In conclusion our results suggest that continued destruction of large trees could affect future numbers of large herbivores in African savannas and better protection of large trees is probably necessary to sustain high animal densities in these ecosystems.     

Dissecting Arabidopsis lateral root development

Recent studies in the model plant Arabidopsis provide new insight into the regulation of root architecture, a key determinant of nutrient- and water-use efficiency in crops. Lateral root (LR) primordia originate from a subset of pericycle founder cells. Sophisticated mass-spectroscopy-based techniques have been used to map the sites of biosynthesis of auxin and its distribution in Arabidopsis seedlings, highlighting the importance of the phytohormone during LR initiation and emergence. Key components of the cell cycle and signal-transduction pathway(s) that promote and attenuate auxin-dependent LR initiation have recently been identified. Additional signals, such as abscisic acid and nitrate, also regulate LR emergence, raising intriguing questions about the cross-talk between their transduction pathways.