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About annbot

Ann Bot is a gestalt entity who works in the office for the Annals of Botany.

Seed dormancy and parasitic plants – this week in Annals of Botany

Acacia saligna Humidity-regulated dormancy onset in the Fabaceae: a conceptual model and its ecological implications for the Australian wattle Acacia saligna
Seed dormancy enhances fitness by preventing seeds from germinating when the probability of seedling survival and recruitment is low. The onset of physical dormancy is sensitive to humidity during ripening; however, the implications of this mechanism for seed bank dynamics have not been quantified. This study proposes a model that describes how humidity-regulated dormancy onset may control the accumulation of a dormant seed bank, and seed experiments are conducted to calibrate the model for an Australian Fabaceae, Acacia saligna. The model is used to investigate the impact of climate on seed dormancy and to forecast the ecological implications of human-induced climate change.

 

Arabinogalactan protein-rich cell walls, paramural deposits and ergastic globules define the hyaline bodies of rhinanthoid Orobanchaceae haustoria
Parasitic plants obtain nutrients from their hosts through organs called haustoria. The hyaline body is a specialized parenchymatous tissue occupying the central parts of haustoria in many Orobanchaceae species. The structure and functions of hyaline bodies are poorly understood despite their apparent necessity for the proper functioning of haustoria. This paper reports a cell wall-focused immunohistochemical study of the hyaline bodies of three species from the ecologically important clade of rhinanthoid Orobanchaceae.

 

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Plants and Climate Change: Complexities and Surprises

Photo: BigStockPhoto.

Photo: BigStockPhoto.

Four decades of intensive research into anthropogenically induced shifts in CO2, precipitation, and temperature evidence important biological impacts on many plant species. As ecologists develop more sophisticated experiments however, many unexpected responses become apparent, suggesting that response to climate change in real world vegetation is more complex than the experiments and models of past decades could be expected to anticipate. Nonetheless, these complexities must be understood if we are to have any hope of predicting the effects of anthropogenic climate change on biological systems.

In this Symposium session at ESA 2014, we focus on surprises in plant responses, highlighting mismatches between theory, modelling, experimental and observational studies. We bring together expertise from multiple levels of study (from individual to ecosystems), using manifold approaches (from experimental to modelling to observational), and from ecological, evolutionary and paleo perspectives. Speakers span a range of career stages, from well-established to just finishing their PhDs and include perspectives from four continents (North America, Africa, Asia and Australia).

By bringing together this diversity of topics, approaches and perspectives, we aim to gain new insights and promote future interdisciplinary research on plant/climate interactions.

We hope you’ll be able to join us in Sacramento.

Camille Parmesan, Marine Institute, University of Plymouth, UK. and
Mick Hanley, School of Biological Sciences, University of Plymouth, UK.

Speakers

13:30 – Richard Primack (University of Boston, USA) Autumn leaf phenology: A search for patterns using 1000 species at four botanical gardens

14:00 – Susan P. Harrison (University of California – Davis, USA) Ecological contingency in the effects of climate change on plant communities: reconciling experimental, historical, interannual, and geographic evidence

14:30 – Kumar P. Mainali (University of Texas at Austin, USA) Complex drivers of population dynamics across treeline: expected and unexpected responses in Himalayan systems

15:10 – Osvaldo Sala (Arizona State University, USA) Lags in the response of ecosystems to directional changes in water availability

15:40 – Tianhua He (Curtin University, Perth, Australia) In situ evolutionary adaptation of Australian plants to climate change

16:10 – Guy Midgley (Stellenbosch University, South Africa) Individualistic species vs. ecosystem responses in under changing climate and CO2 conditions

16:40 Panel Discussion

Annals of Botany Special Issue

The Annals of Botany journal expects to publish work presented at Sacramento as part of a special issue on Plants and Climate Change in the early part of 2015.
The journal welcomes submission of relevant papers from any plant biologist or ecologist for publication in this special issue. Please contact Mick Hanley (mehanley(at)plymouth.ac.uk) for more details.

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Orchid pollen, clover and salinity and Arabidopsis REM – This Week in Annals of Botany

Dactylorhiza maculata Desiccation tolerance, longevity and seed-siring ability of entomophilous pollen from UK native orchid species
Pollinator-limited seed-set in some terrestrial orchids is compensated for by the presence of long-lived flowers. This study tests the hypothesis that pollen from these insect-pollinated orchids should be desiccation tolerant and relatively long lived using four closely related UK terrestrial species; Anacamptis morio, Dactylorhiza fuchsii, D. maculata and Orchis mascula.

 

Heritability and quantitative genetic divergence of serotiny, a fire-persistence plant trait
Although it is well known that fire acts as a selective pressure shaping plant phenotypes, there are no quantitative estimates of the heritability of any trait related to plant persistence under recurrent fires, such as serotiny. In this study, the heritability of serotiny in Pinus halepensis is calculated, and an evaluation is made as to whether fire has left a selection signature on the level of serotiny among populations by comparing the genetic divergence of serotiny with the expected divergence of neutral molecular markers.

 

Leaf hydraulic vulnerability to drought is linked to site water availability across a broad range of species and climates
Vulnerability of the leaf hydraulic pathway to water-stress-induced dysfunction is a key component of drought tolerance in plants and may be important in defining species’ climatic range. However, the generality of the association between leaf hydraulic vulnerability and climate across species and sites remains to be tested.

 

Salinity-mediated cyanogenesis in white clover (Trifolium repens) affects trophic interactions
Increasing soil salinity poses a major plant stress in agro-ecosystems worldwide. Surprisingly little is known about the quantitative effect of elevated salinity on secondary metabolism in many agricultural crops. Such salt-mediated changes in defence-associated compounds may significantly alter the quality of food and forage plants as well as their resistance against pests. In this study, the effects of soil salinity on cyanogenesis in white clover (Trifolium repens), a forage crop of international importance, are analysed.

 

Analysis of the arabidopsis REM gene family predicts functions during flower development
The REM (Reproductive Meristem) gene family of Arabidopsis thaliana is part of the B3 DNA-binding domain superfamily. Despite the fact that several groups have worked on the REM genes for many years, little is known about the function of this transcription factor family. This study aims to identify a set of REM genes involved in flower development and to characterize their function.

 

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This Week in Annals of Botany

Light microscopy images of wood Automatic identification and characterization of radial files in light microscopy images of wood
Analysis of anatomical sections of wood provides important information for understanding the secondary growth and development of plants. This study reports on a new method for the automatic detection and characterization of cell files in wood images obtained by light microscopy.

 

Using virtual 3-D plant architecture to assess fungal pathogen splash dispersal in heterogeneous canopies: a case study with cultivar mixtures and a non-specialized disease causal agent
Recent developments in plant disease management have led to a growing interest in alternative strategies, such as increasing host diversity and decreasing the use of pesticides. Use of cultivar mixtures is one option, allowing the spread of plant epidemics to be slowed down. As dispersal of fungal foliar pathogens over short distances by rain-splash droplets is a major contibutor to the spread of disease, this study focused on modelling the physical mechanisms involved in dispersal of a non-specialized pathogen within heterogeneous canopies of cultivar mixtures, with the aim of optimizing host diversification at the intra-field level.

 

AGO1 controls arabidopsis inflorescence architecture possibly by regulating TFL1 expression
The TERMINAL FLOWER 1 (TFL1) gene is pivotal in the control of inflorescence architecture in arabidopsis. Thus, tfl1 mutants flower early and have a very short inflorescence phase, while TFL1-overexpressing plants have extended vegetative and inflorescence phases, producing many coflorescences. TFL1 is expressed in the shoot meristems, never in the flowers. In the inflorescence apex, TFL1 keeps the floral genes LEAFY (LFY) and APETALA1 (AP1) restricted to the flower, while LFY and AP1 restrict TFL1 to the inflorescence meristem. In spite of the central role of TFL1 in inflorescence architecture, regulation of its expression is poorly understood. This study aims to expand the understanding of inflorescence development by identifying and studying novel TFL1 regulators.

 

Explaining ontogenetic shifts in root-shoot scaling with transient dynamics
Simple models of herbaceous plant growth based on optimal partitioning theory predict, at steady state, an isometric relationship between shoot and root biomass during plant ontogeny, i.e. a constant root–shoot ratio. This prediction has received mixed empirical support, suggesting either that optimal partitioning is too coarse an assumption to model plant biomass allocation, or that additional processes need to be modelled to account for empirical findings within the optimal partitioning framework. In this study, simulations are used to compare quantitatively two potential explanations for observed non-isometric relationships, namely nutrient limitation during the experiments and initial developmental constraints.

 

Callose biosynthesis in arabidopsis with a focus on pathogen response: what we have learned within the last decade
(1,3)-β-Glucan callose is a cell wall polymer that is involved in several fundamental biological processes, ranging from plant development to the response to abiotic and biotic stresses. Despite its importance in maintaining plant integrity and plant defence, knowledge about the regulation of callose biosynthesis at its diverse sites of action within the plant is still limited. Arabidopsis (Arabidopsis thaliana) is one of the best-studied models not only for general plant defence responses but also for the regulation of pathogen-induced callose biosynthesis. This article summarizes what is known about the regulation of callose synthase activity as well as what has been discussed with regard to this topic within the last decade based on results derived from new techniques and available mutant lines, focusing on the progress that has been made in understanding the regulation of callose biosynthesis in response to pathogen attack.

 

Linking ecophysiological modelling with quantitative genetics to support marker-assisted crop design for improved yields of rice under drought stress
Genetic markers can be used in combination with ecophysiological crop models to predict the performance of genotypes. Crop models can estimate the contribution of individual markers to crop performance in given environments. This study explores the use of crop models to design markers and virtual ideotypes for improving yields of rice (Oryza sativa) under drought stress.

 

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Impact of climate on plant growth and flower formation – This Week in Annals of Botany

Xylem Impact of warming and drought on carbon balance related to wood formation in black spruce
Wood formation in trees represents a carbon sink that can be modified in the case of stress. The way carbon metabolism constrains growth during stress periods (high temperature and water deficit) is now under debate. In this study, the amounts of non-structural carbohydrates for xylogenesis in black spruce saplings were assessed under high temperature and drought in order to determine the role of sugar mobilization for osmotic purposes and its consequences for secondary growth. Plant water status during wood formation can influence the materials available for growth in the cambium and xylem.

 

Relative growth rate variation of evergreen and deciduous savanna tree species is driven by different traits
Plant relative growth rate depends on biomass allocation to leaves (leaf mass fraction, efficient construction of leaf surface area (specific leaf area) and biomass growth per unit leaf area (net assimilation rate). This paper shows that trade-offs between investment in carbohydrate reserves and growth occur only among deciduous species, leading to differences in relative contribution made by the underlying components of relative growth rate between the leaf habit groups. The results suggest that differences in drivers of relative growth rate occur among savanna species because these have different selected strategies for coping with fire disturbance in savannas.

 

DEF- and GLO-like proteins may have lost most of their interaction partners during angiosperm evolution
DEFICIENS (DEF)- and GLOBOSA (GLO)-like proteins constitute two groups of floral homeotic transcription factors that were already present in the most recent common ancestor of angiosperms. Together they specify the identity of petals and stamens in flowering plants. This paper strengthens the hypothesis that a reduction in the number of interaction partners of DEF- and GLO-like proteins, with DEF–GLO heterodimers remaining the only DNA-binding dimers in core eudicots, contributing to developmental robustness, canalization of flower development and the diversification of angiosperms.

 

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What inspired you to do plant science? #epso2014

This week guest author Charlie Haynes is AoB Blog’s roving reporter at the EPSO/FESPB plant biology Europe conference.

At the FESPB/EPSO plant biology conference in Dublin I asked some of the delegates what inspired them to work in plant science, botany and ecology. Here are just a few of their answers:

 

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AoB Interviews: Hans Lambers on soil phosphate acquisition in impoverished soil

This week guest author Charlie Haynes is AoB Blog’s roving reporter at the EPSO/FESPB plant biology Europe conference.

 

Hans Lambers Hans Lambers is the Winthrop Professor at the University of Western Australia. He competed his PhD in 1979 at the University of Groningen in the Netherlands and since then worked at Melbourne University, Australian National University and Utrecht University. His research focuses on mineral nutrition of native Australian plants and crop and pasture legumes. He very kindly agreed to talk to me about some of the challenges of soil phosphate impoverishment.

Why is phosphate impoverishment so significant?
It’s of less importance in Europe which imports food and animal feed from parts of the world where phosphates passing the problem on. There it is an issue of excess of phosphate, dumped on the land and ending up in waterways. Europe could stop fertilising now and still have crops for the next 20 years. But when you go to other parts of the world; Australia, South America, Africa and South East Asia, phosphate insecurity is a real issue. This may be because the amount in the soil is too low for effective crop production, or it may be that it is there but it’s not readily available. So it’s an issue for crop production and thus food security. What we can do though is instead grow plants that can use that phosphorus in the soil much more effectively. There is a tremendous opportunity.

How does this limit these countries in what they can grow and the yields they can produce?
In Africa phosphorus is the key limiting factor, even some the driest areas of Saharan Africa. People who worked in barchenener discovered that simply by adding phosphorus you could get a higher yield. The dry soil significantly reduces the mobility of phosphorus in the soil and it becomes a significant limiting when you have a dry soil (Lambers H, Raven JA, Shaver GR, Smith SE. (2008) Plant nutrient-acquisition strategies change with soil age. Trends in Ecology and Evolution 23: 95-103).

So why do these communities not buy fertiliser to increase their yield?
Fertiliser is very expensive for these groups. It has to travel vast distances to the harvest, and these groups simply don’t have the money for it. So there instead we are working towards crops that are more efficient at using the existing phosphorus or are better at getting it out of the soil. This is however a bit of a risky business – if you have soils that are very nutrient poor to begin with then plants that extract it more effectively will make the soil even more phosphorus deficient. Whatever you take out of soil have to replace in order to be sustainable.

What makes phosphates accessible?
Phosphates in soil are readily available at a neutral pH. Calcareous soils with their more alkali pH lock phosphates up in calcium complexes. The phosphate is there but not readily available to crops. More acidic soils also lock up phosphates – but this time not in calcium complexes but instead, as complexes of oxides and hydroxides of iron and aluminium. Chile has very acidic soil, with a pH of almost 4, and lots of these metal oxides and hydroxides so all the phosphate isn’t readily available. However the plants have special adaptations that allow them to access it in these conditions.

What are these adaptions?
These plants have a special structure which works in combination with the plant biochemistry. What they produce is massive quantities of carboxylates. These are molecules with a negative charge – like phosphate. These exchange for one another, releasing the phosphate ions into the soil solution, whilst carboxylates anion take the place of phosphate in the soil. You’re effectively mining the phosphate that is in the soil out of it’s tight bindings. It’s then in the solution and anybody can take it up (Lambers H, Bishop JG, Hopper SD, Laliberté E, Zúñiga-Feest A. (2012) Phosphorus-mobilization ecosystem engineering: the roles of cluster roots and carboxylate exudation in young P-limited ecosystems. Annals of Botany 110: 329-348).

Could this be put into another crop either by breeding or genetic modification?
I would take one step back and ask ‘what crops do we have now at already can do that’? White Lupin is an excellent example and there are a few other Lupin species that do exactly the same. There are also some Lupin species that don’t have these wonderful structures but instead something close to it, and some without a structure at all that still release carboxylates. So we actually already have a lot of species that can already played this trick. Rather than engineer this in Soybean, it’s important to get a thorough understanding of the technology. Understanding is and farming it in crops with the gene is an obvious first stage. We already have crops with the this ability in lupins – which are much better than wheat and barley at this stage. I don’t think it’s impossible but it’s important to take it one step at a time (Lambers H, Clements JC, Nelson MN. (2013) How a phosphorus-acquisition strategy based on carboxylate exudation powers the success and agronomic potential of lupines (Lupinus, Fabaceae). American Journal of Botany 100: 263-288).

So are some parts of the world focusing on the wrong crops for their soil type and climate?
Yes absolutely! In chile they used to grow Andean Lupin. When the Spanish invaded they forbade the natives from growing these lupins as they weren’t Spanish crops. The natives switching to foreign crops is a daft idea when they already had a crop suited to their environment! Quinoa is an example of another crop where this happened, and the Spanish stopped that. They arguably had better crops than the foreign spanish ones then introduced. One thing one can do though is intercropping. This is where you grow plants concurrently interspersed between one another. If you want to grow wheat, it cannot grow particularly well in some South American environments. If you intercrop it with Lupin, can mobilise that phosphorus and the neighbours can benefit from that. You can also do crop rotation. A group in Germany has actually done this, working with rotations of soybean and maize. Maize is not so good at accessing phosphate, soybean – depending on the cultivar you use, is. The good soybean cultivars show a real benefit for the next crop – a phosphorus benefit. You can grow them at the same time or you can grow them in rotation to access this phosphate. Both of these techniques have tremendous benefits.

What stops people in phosphorus poor environments from doing this already?
That’s an interesting question. If you go to china, intercropping has been done for hundreds of years and you can demonstrate that with the right intercropping combinations you can have a 40-50% higher yield – which is pretty impressive! A British or Irish farmer with an increase in yield of that kind of level would be ecstatic! So the Chinese already have done that, and Europe is exploring it. I’m certain it could be done in other parts of the world, but it’s not happening on a large scale and that’s because a lack of education. It’s important to educate local farmers about this from Africa to Australia! I’m working with a group in Germany Andreas Burgutts, who is screening sorghums for better phosphate accessibility, using leaf manganese levels as a marker. These are taken up by the plant in the same as as phosphates and so used as a marker. Work like this requires going to Africa, and selecting the right cultivar for the conditions there, not in our lab field. This is about doing research and then making use this research reaches farmers, and doesn’t stay in a scientist’s ivory tower. Work needs to be done and go beyond journals, into places where we can make a difference,

Who else is working on taking this knowledge into the field?
I had a visit from someone from ICRISAT. They are based in India and work on major crop draught and salinity. They are now keen to work on phosphorus, and they had heard of my work and were interested in developing something together. These big international institutes have the links with the grassroots farming communities in the parts of the world where you can truly make a difference. I may be able to do high end science but without the connections I’m not able to have much of a real world impact.

Who else is involved?
The big international institutes are doing good work, IRRI in the Philippines, ICARDA in Aleppo and ICRISAT in Hyderabad. These large international institutes aren’t just interested in the science, but also applying it, and I think that is really important.

Do these plants have potential in any other key areas?
Yes, for instance where you have soil contaminated with heavy metals you could use them in the process of photoremediation. Here plants as used for their ability to remove heavy metals an ‘clean’ soil. There are areas in Belgium that have been heavily polluted with zinc or copper. Chemical or physical cleaning of this soil is almost impossible. You need a species that accumulates these metals to a very high concentration, but is also a fast grower, producing a lot of biomass, or else the process takes along time. There is serious potential in this. In addition to this these plants can be used in phytomining or prospecting, accumulating small amounts of metals that act as an indicator for a larger deposit of metal in the earth. This can act as a pretty good indicator of gold and some other metals to allow groups to commence mining.
Hans’ book “Plant Life on the Sandplains in Southwest Australia, a Global Biodiversity Hotspot” will be out in September, and is now available online at the UWA Publishing website.

 

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Why Botany is Important #epso2014

Society of Biology calls for investment in plant science This week guest author Charlie Haynes is AoB Blog’s roving reporter at the EPSO/FESPB plant biology Europe conference. This post is his pre-conference manifesto.

 

On the first day of the EPSO/FESPB plant biology Europe conference it’s worth considering why botany is important.

Like many others whilst studying GCSE and A Level biology I found the botanical themed part of the syllabus dull and uninteresting. I arrived at university to find myself surrounded by those with similar experiences in their schools. Not one person I met during my first year of Biological Sciences at Leicester said they wanted to be a professional botanist. Luckily I turned up to all of my lectures and found myself interested and maybe even enjoying some of the botany and ecology modules that I was initially less than thrilled about taking. But there are serious emerging issues in plant science and ecology that need more talent.

  • A burgeoning world population needs ever greater crop yields as people become increasingly affluent and demand a higher quality and quantity of food produce.
  • Climate change is increasing the incidence of severe weather conditions such as droughts and heavy rains. Some climate models show that with a temperature rise of 2 degrees Celsius by 2050 will lower wheat yields by an average of 50%.
  • Despite large scale agricultural enterprises, an estimated 50% of world food production is from small small farmers. Many of these are subsistence farmers. The average Vietnamese farm is approximately 340 times smaller than the average US farm. New innovative strategies need to help these farmers use this space effectively.
  • Disease can still strike harvests dead in their tracks destroying livelihoods and causing skyrocketing food prices.
  • With more individuals demanding a western style lifestyle, the need for fresh water is also climbing. A potato has a water footprint of 25 litres. A hamburger has a water footprint of an estimated 2400 litres. But we live on a planet where only 2.5% of all water is fresh water and much of this is trapped at the polar ice caps. Developing plants capable of reducing their water footprint is vital in some chronically dry regions.
  • Nutritional deficits and diseases account for millions of deaths and over 2 billion are malnourished. Not only does the total number of calories produced need to increase, but there also needs to be an increase in global dietary variance and quality.

Have I missed anything else blindingly obviously that screams a need for plant science in the 21st century?

If so please let me know in the comments below, I would love to hear from you!

 

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This week in Annals of Botany – bumper plant science edition!

Sapria himalayana Holoparasitic Rafflesiaceae possess the most reduced endophytes and yet give rise to the world’s largest flowers
Species in the parasitic plant family Rafflesiaceae exhibit one of the most highly modified vegetative bodies in flowering plants. Apart from the flower shoot and associated bracts, the parasite is a mycelium-like endophyte living inside their grapevine hosts. This study provides a comprehensive treatment of the endophytic vegetative body for all three genera of Rafflesiaceae (Rafflesia, Rhizanthes and Sapria), and reports on the cytology and development of the endophyte, including its structural connection to the host, shedding light on the poorly understood nature of this symbiosis.

 

Plant traits and ecosystem effects of clonality: a new research agenda
Clonal plants spread laterally by spacers between their ramets (shoot–root units); these spacers can transport and store resources. While much is known about how clonality promotes plant fitness, we know little about how different clonal plants influence ecosystem functions related to carbon, nutrient and water cycling. This review gives some concrete pointers as to how to implement this new research agenda through a combination of (1) standardized screening of predominant species in ecosystems for clonal response traits and for effect traits related to carbon, nutrient and water cycling; (2) analysing the overlap between variation in these response traits and effect traits across species; (3) linking spatial and temporal patterns of clonal species in the field to those for soil properties related to carbon, nutrient and water stocks and dynamics; and (4) studying the effects of biotic interactions and feedbacks between resource heterogeneity and clonality. Linking these to environmental changes may help us to better understand and predict the role of clonal plants in modulating impacts of climate change and human activities on ecosystem functions.

 

Selenium addition alters mercury uptake, bioavailability in the rhizosphere and root anatomy of rice
Mercury is an extremely toxic pollutant, especially in the form of methylmercury, whereas selenium is an essential trace element in the human diet. This study aimed to ascertain whether addition of selenium can produce rice with enriched selenium and lowered mercury content when growing in mercury-contaminated paddy fields and, if so, to determine the possible mechanisms behind these effects.

 

Patch size and distance: modelling habitat structure from the perspective of clonal growth
In a patchy habitat, parents and their offspring can be under different environmental conditions, for example in terms of light/shade or high/low nutrient concentration. This study considers the spatial structure of patchy habitats from the perspective of plants that forage for resources by clonal growth. Modelling is used in order to compare two basic strategies, which differ in the response of the plant to a patch boundary. The results suggest that the degree of physiological integration between a parent and an offspring ramet is important even across a very short distance because it can strongly influence the efficiency of foraging.

 

Variable response of three white clover ecotypes to soil flooding by seawater
Despite concerns about the impact of rising sea levels and storm surge events on coastal ecosystems, there is remarkably little information on the response of terrestrial coastal plant species to seawater inundation. This study examines the responses of a glycophyte (white clover, Trifolium repens) to short-duration soil flooding by seawater and recovery following leaching of salts and suggests that selection for tolerant ecotypes is possible should the predicted increase in frequency of storm surge flooding events occur.

 

Changes in tracheid and ray traits in fire scars of North American conifers and their ecophysiological implications
Fire scars have been widely used as proxies for the reconstruction of fire history. However, little is known about the impact of fire injury on wood anatomy. This study investigates changes in tracheid and ray traits in fire scars of Douglas fir (Pseudotsuga menziesii), western larch (Larix occidentalis) and ponderosa pine (Pinus ponderosa), and discusses their ecophysiological implications for tree recovery from fire.

 

Morphological and physiological divergences within holm oak support the existence of different ecotypes depending on climatic dryness
Previous studies show contradictory findings about the functional convergence within the Mediterranean woody flora. This study evaluates the variability of functional traits within holm oak (Quercus ilex) to elucidate whether provenances corresponding to different morphotypes represent different ecotypes locally adapted to the prevaling stress levels. This is the first time that the combined use of morphological and physiological traits has provided support for the concept of these two holm oak morphotypes being regarded as two different species.

 

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