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Cones are slightly deflexed on short stout stalks, symmetrical, hard, heavy, ovoid-conic, 7- cm long, cm broad when closed, ripening rich glossy chestnut-brown in April two years after pollination, and opening the same summer or up to 10 years later, to cm broad.

Balearic Islands. Valencia , and Tunisia. Bannister and Neuner describe it as hardy to Zone 8 cold hardiness limit between However, a common-garden experiment by Prada et al. The coastal populations displayed the opposite behavior, while the continental Moroccan population presented a unique combination of traits. We confirmed trade-offs between cold-hardiness and growth at the population level, but not within populations. The largest recorded specimen appears to have been 4.

The tallest is stated to be There is a record of a The tallest ever recorded, according to a chief technician of the Forests National Office France , was a 42 m tall tree in the forest of the Landes S. As of Half of these studies have looked at the species' utility in reconstructing past climate variation, and most of the others have looked at ecological problems such as stand development, productivity, and the impact of insect defoliators.

The largest man-made forest in the world, the , ha Les Landes on the Atlantic coast of SW France, is planted almost entirely with this species. This was originally planted from onward not for timber, but for land reclamation, with a huge area of shifting sand dunes threatening fertile farmland futher inland Frankis, M. The species has been widely planted in other parts of the world with a Mediterranean climate, and is now naturalised in South Africa and elsewhere Price et al.

Aiton, W. Hortus Kewensis vol. Available: www. Gaussen, H. Heywood, and A. In: T. Tutin et al. Greuter, W. Burdet, and G. Checklist - A critical inventory of vascular plants of the circum-mediterranean countries , Vol. Life-history correlations with seasonal cold hardiness in maritime pine.

American Journal of Botany 12 Elwes and Henry at the Biodiversity Heritage Library. This series of volumes, privately printed, provides some of the most engaging descriptions of conifers ever published. It is not very long-lived in cultivation[11, ], though trees have lived longer than years in Britain. Widely cultivated in Europe for the resin obtained from the trunk and for woodpulp, it is the most widely used European species[64, ].

Plants are strongly outbreeding, self-fertilized seed usually grows poorly[]. They hybridize freely with other members of this genus[]. Leaf secretions inhibit the germination of seeds, thereby reducing the amount of plants that can grow beneath the trees[18]. Plants in this genus are notably susceptible to honey fungus[]. Celsius Fahrenheit:.

Plants For A Future have a number of books available in paperback and digital form. Our new book to be released soon is Edible Shrubs. It is best to sow the seed in individual pots in a cold frame as soon as it is ripe if this is possible otherwise in late winter. Plant seedlings out into their permanent positions as soon as possible and protect them for their first winter or two[11]. Plants have a very sparse root system and the sooner they are planted into their permanent positions the better they will grow[K].

Trees should be planted into their permanent positions when they are quite small, between 30 and 90cm[]. We actually plant them out when they are about 5 - 10cm tall. So long as they are given a very good weed-excluding mulch they establish very well[K]. Larger trees will check badly and hardly put on any growth for several years.

This also badly affects root development and wind resistance[]. This method only works when taken from very young trees less than 10 years old. Use single leaf fascicles with the base of the short shoot. Disbudding the shoots some weeks before taking the cuttings can help. Cuttings are normally slow to grow away[81]. Right plant wrong place. We are currently updating this section. Author Aiton. For a list of references used on this page please go here.

If you have important information about this plant that may help other users please add a comment or link below. Only comments or links that are felt to be directly relevant to a plant will be included. If you have questions about a plant please use the Forum on this website as we do not have the resources to answer questions ourselves.

To leave a comment please Register or login here All comments need to be approved so will not appear immediately. You can unsubscribe at anytime. Follow Us:. Pinus pinaster - Aiton. Avoid if allergies. Avoid internally if suffering from asthma or bronchitis. The astringent taste may cause stomach discomfort []. Habitats Coastal sands and dunes, it is also found on mountains inland.

Range Europe - W. Naturalized in Britain in some parts of the New Forest[81]. Translate this page:. Pinus pinaster is an evergreen Tree growing to 20 m 65ft by 7 m 23ft at a fast rate. See above for USDA hardiness. It is hardy to UK zone 8. It is in leaf all year, in flower in May, and the seeds ripen in April.

The species is monoecious individual flowers are either male or female, but both sexes can be found on the same plant and is pollinated by Wind. The plant is not self-fertile. Suitable for: light sandy and medium loamy soils, prefers well-drained soil and can grow in nutritionally poor soil. Suitable pH: mildly acid and neutral soils.

It cannot grow in the shade. It prefers dry or moist soil and can tolerate drought. The plant can tolerate maritime exposure. Special Uses Carbon Farming. Carbon Farming Industrial Crop: Hydrocarbon Materials, chemicals and energy include bioplastics, rubber, biomass products gasoline, jet fuel, diesel, butane, propane, biogas. Plants are usually resprouting plants and saps.

Management: Standard Plants grow to their standard height. Harvest fruit, seeds, or other products.

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At each sampling point physiological measurements were performed on symptomatic plants and controls of the corresponding pine species and plant material was harvested for further analysis. Physiological measurements comprised needle gas-exchange related parameters, relative water content RWC , and electrolyte leakage, and water potential. Fungal-infected plant material and experimental material that contacted with the fungus including soil and containers were adequately decontaminated by autoclaving.

Relative internal stem lesion length was determined in 5—7 cm longitudinal stem cuts of six biological replicates per treatment. This was calculated as a proportion of the total stem length. From each of the six replicates per treatment, five needles were collected, fresh weight FW was recorded, and needles were transferred to tubes with distilled water. Controlled conditions were maintained inside the chamber: ambient CO 2 concentration Data were recorded when parameters remained stable.

Six biological replicates per treatment were measured. Three needles from the vegetative growing year per plant were collected into 15 mL Falcon tubes, cut into 1 cm pieces, immersed in 10 mL of Milli-Q water and incubated for 12 h room temperature, rpm. Conductivity was measured C exp , samples were autoclaved 20 min. Initial Milli-Q water conductivity C i was recorded, as well as after autoclaving C ii.

Six biological replicates per treatment were considered. To determine pigment concentration six biological replicates per treatment were considered. Total chlorophyll concentration was calculated following authors recommendations. The homogenate was immersed in boiling water for 1. Anthocyanin concentration was determined according to the equation proposed by Close et al. The acidified water extract was partitioned twice against 1.

The organic upper layer was recovered and vacuum evaporated in a centrifugal concentrator Speed Vac, Jouan, Saint Herblain Cedex, France. The resulting solution was passed through 0. Analytes were separated by reversed-phase Nucleodur C18, 1. Hormones were quantified with a Quattro LC triple quadrupole mass spectrometer Micromass, Manchester, United Kingdom connected online to the output of the column through an orthogonal Z-spray electrospray ion source. Quantitation of hormones was achieved using a standard curve.

Six biological replicates per treatment were analysed. RNA was extracted from 50 mg of pine needles of six biological replicates per treatment using the method described by Valledor et al. For each treatment, two cDNA pools of three biological replicates each were used. The relative abundance of transcripts of a set of target genes related to plant primary metabolism and pathogen response was assessed by Real-Time qPCR Table 1. Primer pairs were designed using Primer3 v.

Three analytical replicates of the two cDNA pools per treatment and pine species were performed. The corresponding reference genes were included in every plate. Gene expression results were obtained following the recommendations of Hellemans et al. Amplicon band size was confirmed by agarose gel for every gene. Biological variations were controlled by analysing quality control QC standards by fatty acid methyl esters internal standard markers and a QC standard solution of 41 pure reference compounds i.

Primary metabolites were annotated using TagFinder Luedemann et al. The relative abundance of primary metabolite levels was normalised to the internal standard ribitol and the dry weight of the samples. Physiological, hormone and gene expression data were analysed using SigmaPlot v. Asterisks indicate significant differences between non-inoculated controls C and plants inoculated with F.

Gene expression levels were used as predictor matrix for metabolite and physiological responses. Models were tuned based on total Q2 threshold for component consideration was established at 0. The same symptoms were registered 8 d.

Figure 1A ; ca. Pinus pinea remained asymptomatic for longer, revealing slight disease symptoms 64 d. Only P. Figure 1. Progression of pitch canker disease in Pinus. A Time course of the percentage of plants inoculated with F. Images representative of the symptoms at each sampling point are presented. Days post-infection d. B Stem relative internal necrosis length of non-inoculated controls black bars and of plants inoculated with F.

C Stem internal necrosis representative of non-inoculated controls left column and plants inoculated with F. Physiological and biochemical experiments were undertaken at 10, 17 and 64 d. Pinus radiata water potential was significantly decreased after F. While a sharper decrease was observed in P. Fusarium circinatum inoculation significantly decreased RWC in P. Figure 2. Plant water status of non-inoculated controls black bars and plants inoculated with F. A Midday water potential.

B Relative water content. Inoculation with F. In contrast, inoculated P. After F. This was associated with a net increase in needle CO 2 concentration suggesting that decreased assimilation rates were at least partly caused by biochemical of photochemical limitations.

Figure 3. Needle gas exchange-related parameters of non-inoculated controls black bars and plants inoculated with F. A Stomatal conductance. B Transpiration rate. C Net CO 2 assimilation rate. D Sub-stomatal CO 2 concentration. Pinus pinea chlorophyll concentration remained unchanged.

Inoculated P. Table 2. Total chlorophyll and anthocyanin concentration, and electrolyte leakage of non-inoculated control plants C and plants inoculated with F. Inoculation of P. On the other hand, F. In contrast, no change was observed in inoculated P. Figure 4. Hormones concentration of non-inoculated controls black bars and plants inoculated with F. A Abscisic acid.

B Salicylic acid. C Jasmonic acid. No significant changes were observed after F. RuBisCO transcript abundance was significantly reduced in the presence of F. On the contrary, F. PDC and pal transcripts were also significantly increased in inoculated P. Transcript abundance of pr5 was only significantly altered in P.

Transcripts encoding SnRK2. Figure 5. Relative abundance of primary metabolism- and pathogenesis-related genes in Pinus inoculated with F. B Cell wall invertase cwINV. D Pyruvate decarboxylase PDC. E Glycolate oxidase GOX. F Sucrose non-fermenting 1-related protein kinase 2. G Phenylalanine ammonia lyase pal. H Thaumatin-like protein pr5. I Chitinase pr3. The abundance of transcripts in plants inoculated with F. Asterisks indicate significant differences between non-inoculated controls and plants inoculated with F.

Changes in the relative abundance of primary metabolites from all chemical groups with the exception of OA were observed following inoculation with F. Figure 6. Primary metabolite changes occurring in Pinus inoculated with F. Relative values are normalised to the internal standard ribitol and dry weight DW of the samples. Grey-colour square represents not detected n.

Significant increases occurred in several AA in P. The relative abundance of some of these metabolites was also enhanced by the presence of F. Proline Pro levels were also significantly increased by over fold upon F. In inoculated P. Glucose Glc and Fru levels were significantly increased following pathogen infection in P. Sugar alcohols content was altered by PPC: a significant increase was registered for glycerol in inoculated P. Upon F. Additionally, a significant decrease of dehydroascorbate was detected only in P.

A clear separation between control and inoculated samples was found for P. For P. The positioning of inoculated P. Together with this, relative necrosis, Ci, ABA and total chlorophyll are placed in the positive side of the y -axis Figure 7B. On the negative part of the x -axis we find the genes analysed, water relations, A, E, gs, and SA, influencing the overlap observed in P. Figure 7. Integrated data analysis. A Sparse partial least squares sPLS regression analysis of the complete dataset of physiological, hormonal, gene expression and primary metabolism alterations occurring in Pinus inoculated with F.

First two components are plotted in the graph. B Correlation circle plot of the components represented in the sPLS regression analysis. Gene expression levels blue were used as predictor matrix for metabolite and physiological responses orange. The timing and intensity of disease symptoms observed after F. The overview of our data through sPLS analysis also revealed different profiles of response upon pathogen inoculation: P.

Under this water deprivation-like scenario P. The accumulation of ABA upon F. In accordance with a greater water limiting-like status, only inoculated P. Besides conferring tolerance to soybean cyst nematode Matthews et al. The enhanced resilience of P. The increase of chlorophyll concentration in the susceptible Pinus species is at odds with previous studies indicating that photosynthetic gene expression including those associated with pigment biosynthesis is downregulated in a range of species upon pathogen attack Bilgin et al.

In fact, net CO 2 assimilation rate was impaired in inoculated P. Although this is in part explained by reduced stomatal conductance, the accumulation of intercellular CO 2 suggests that also biochemical limitations impaired photosynthesis in these species, as shown by the down-regulation of RuBisCO after F. The down-regulation of photosynthetic genes under pathogen attack is known to be regulated by source-to-sink tissue transformation through cleavage of sucrose into Fru and Glc by cwINV Berger et al.

Compounds from the OPP are involved in the biosynthesis of amino acids AA , such as Trp Maeda and Dudareva, , which accumulated upon pathogen inoculation in the susceptible Pinus species. This may be a means to induce the production of secondary metabolites involved in defence response, as reported in rice infected by Bipolaris oryzae Ishihara et al. However, besides being a common defence response against pathogens, the general accumulation of other AA in P. The decline in P. Phenylalanine plays a key role in linking plant primary and secondary metabolism.

It is used by pal for the synthesis of several compounds crucial to determine plant survival under stressful scenarios Pascual et al. Although pal up-regulation has been associated with F. In fact, the unchanged expression of pr3 and pr5 in P. The overexpression of pr3 in P. The decrease of dehydroascorbate DHA in inoculated P.

Vanacker et al. Moreover, Chen and Gallie demonstrated that DHAR overexpression in guard cells resulted in an increased Asc redox state, reducing H 2 O 2 levels and promoting stomata opening and transpiration. Glycerol accumulation in P.

This response was intensified according to the increasing level of species tolerance, which may indicate species-specific metabolic shifts involved in conferring tolerance to F. In accordance, Tadege et al. PDC converts pyruvate into acetaldehyde, which can enter ethanolic fermentation or the pyruvate dehydrogenase bypass Bolton, The later was involved in Lr34 -mediated wheat resistance to Puccinia triticina but it was not maintained in later stages of infection, explaining the partial resistance observed Bolton et al.

The up-regulation of GOX in inoculated P. Therefore, it is more likely that GOX regulation is related to the lower induction of PDC , which is conceivably associated with increased needle CO 2 even when stomata are closed.

Furthermore, although GOX is known to activate SA-mediated defence responses by Pro and its derivates, no significant changes were found in SA levels for every species. Rather, an increase in JA concentration was observed for inoculated P.

A contrasting response was found for P. Interestingly, SA endogenous levels in controls increase according to the tolerance to F. A significantly higher basal level of SA was also found in an Eucalyptus grandis clone resistant to Chrysoporthe austroafricana in comparison with a susceptible clone Mangwanda et al. This may also be a potential basal mechanism of resistance against F.

Our work provides new insights into the physiological changes associated with pathogenesis and plant response in the Pinus — F. By using an integrated approach, we were able to explore this pathosystem at different cellular levels and we link molecular changes to physiological traits with a focus on primary plant metabolism.

In general P. For the resistant P. These results provide interesting avenues for future research to unveil Pinus—F. Once we understand this relationship we will have better tools to fight PPC. AG-C designed and supervised hormones quantification. JA performed hormone quantification, gene expression experiments, and analysed the data. CA and AMR performed primary metabolite analysis. LV conducted the multivariate analysis. JA wrote the manuscript. All authors discussed the data and reviewed the manuscript.

The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest. We thank Pedro Monteiro and Marta Pitarch for the assistance in the experiments, physiological and hormonal characterisation. Altschul, S. Basic local alignment search tool. Berbegal, M. Evidence for multiple introductions and clonality in Spanish populations of Fusarium circinatum.

Phytopathology , — Berger, S. Plant physiology meets phytopathology: plant primary metabolism and plant-pathogen interactions. Bilgin, D. Biotic stress globally downregulates photosynthesis genes. Plant Cell Environ. Bolton, M. Primary metabolism and plant defense-fuel for the fire. Plant Microbe Interact. Lr34 -mediated leaf rust resistance in wheat: transcript profiling reveals a high energetic demand supported by transient recruitment of multiple metabolic pathways.

First report of pitch canker on pines caused by Fusarium circinatum in Portugal. Plant Dis. Carrasco, A. Expression profiling in Pinus radiata infected with Fusarium circinatum. Tree Genet. Genomes Cerqueira, A. Phosphite shifts physiological and hormonal profile of monterey pine and delays Fusarium circinatum progression. Plant Physiol. Chen, Z. The ascorbic acid redox state controls guard cell signaling and stomatal movement.

Plant Cell 16, — The bark is orange-red, thick, and deeply fissured at the base of the trunk, somewhat thinner in the upper crown. The maritime pine features the longest and most robust needles of all European pine species.

Maritime pine is closely related to Turkish pine , Canary Island pine , and Aleppo pine , which all share many features with it. It is a relatively non-variable species, with constant morphology over the entire range. Its range is in the western Mediterranean Basin , extending from Portugal and Northern Spain especially in Galicia to southern and Western France , east to western Italy, Croatia and south to northern Tunisia , Algeria and northern Morocco. It generally occurs at low to moderate altitudes, mostly from sea level to m 2, ft , but up to 2, m 6, ft in the south of its range in Morocco.

The high degree of fragmentation in the current natural distribution is caused by two factors: the discontinuity and altitude of the mountain ranges causing isolation of even close populations, and human activity. Pinus pinaster is a popular topic in ecology because of its problematic growth and spread in South Africa for the past years after being imported into the region at the end of the 17th century — Fynbos vegetation is a fire-prone shrubland vegetation that is found in the southern and southwest cape of South Africa.

It is found in greater abundance close to watercourses. The species favors acidic soils with medium to high-density vegetation, [6] but it can also grow in basic soils and even in sandy and poor soils, where only few commercial species can grow. Pinus pinaster is a diagnostic species of the vegetation class Pinetea halepensis. Larvae of the moth Dioryctria sylvestrella feed on this pine.

Their boring activity causes large quantities of resin to flow from the wounds which weakens the tree and allows fungi and other pathogens to gain entry. Pinus pinaster is a successful invasive species in South Africa. One of the results of its invasion in South Africa is a decrease in the biodiversity of the native environment. Invasive species occupy habitats of native species often forcing them to extinction or endangerment. In addition, depending on the regions P. This depletes the resources available for other species in the environment.

In the Mpumalanga Province , 6 streams completely dried up 12 years after grasslands were replaced with pines. Once again there was a positive effect when some of the pines were removed and agreeable range grasses were planted. The grazing conditions for the sheep of the area were greatly improved when the P.

The invasion of P. Pinus pinaster is particularly successful in regions with fynbos vegetation because it is adapted to high-intensity fires, thus allowing it to outcompete other species that are not as well adapted to high-intensity fires. In areas of fire-prone shrubland, the cones of P.

This adaptation increases the competitive ability of P. When the height of P. Several other characteristics contribute to their success in the regions they have invaded, including their ability to grow rapidly and to produce small seeds with large wings. Their ability to grow quickly with short juvenile periods allows them to outcompete many native species while their small seeds aids in their dispersal.

The small seeds with large wings are beneficial for wind dispersal, which is the key to reaching new areas in regions with fynbos vegetation. For this reason, the small seed, low seed wing loading, and high winds found in mountainous regions all combine to provide a favorable situation for the dispersal of P. Its dispersal ability is one of the key factors that have allowed P.

In addition to being an efficient disperser, P. According to an experiment done in Spain , the resin canal density was twice as high in the P. Since P. The resins make the P. In other words, P. These characteristics enhance the ability of P. Insects and mites that feed on the seeds and cones of P. An insect or mite that acts as an ideal biological control should have a high reproductive rate and be host-specific , meaning that it preys specifically on P.

The life cycle of the predator should also match that of its specific host. Two key characteristics the predator should also exhibit are self-limitation and the ability to survive in the presence of a declining prey population. Controlling the spread of P. The main advantage to using this mite to control the population of P. Pinaster is its specificity to P.

Another possible option is Pissodes validirostris , a cone-feeding weevil that lays eggs in developing cones. When the larvae hatch, they feed on the growing seed tissue, preventing P. Although the adults feed on the trees as well, they do not do any damage to the seeds and only feed on the shoots of the tree, so they do not appear to negatively impact the growth of the trees.

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