Reduced height (Rht) and photoperiod insensitivity (Ppd) allele associations with establishment and early growth of wheat in contrasting production systems

Near isogenic lines (NILs) varying for genes for reduced height (Rht) and photoperiod insensitivity (Ppd-D1a) in a cv. Mercia background (rht (tall), Rht-B1b, Rht-D1b, Rht-B1c, Rht8c + Ppd-D1a, Rht-D1c, Rht12) were compared at one field site but within contrasting ('organic' vs. 'conventional') rotational and agronomic contexts, in each of 3 years. In the final year, further NILs (rht (tall), Rht-B1b, Rht-D1b, Rht-B1c, Rht-B1b + Rht-D1b, Rht-D1b + Rht-B1c) in both Maris Huntsman and Maris Widgeon backgrounds were added together with 64 lines of a doubled haploid (DH) population [Savannah (Rht-D1b) x Renesansa (Rht-8c + Ppd-D1a)]. Assessments included laboratory tests of germination and coleoptile length, and various field measurements of crop growth between emergence and pre jointing [plant population, tillering, leaf length, ground cover (GC), interception of photosynthetically active radiation (PAR), crop dry matter (DM) and nitrogen accumulation (N), far red: red reflectance ratio (FR:R), crop height, and weed dry matter]. All of the dwarfing alleles except Rht12 in the Mercia background and Rht8c in the DHs were associated with reduced coleoptile length. Most of the dwarfing alleles (depending on background) reduced seed viability. Severe dwarfing alleles (Rht-B1c, Rht-D1c and Rht12) were routinely associated with fewer plant numbers and reduced early crop growth (GC, PAR, DM, N, FR:R), and in 1 year, increased weed DM. In the Mercia background and the DHs the semi-dwarfing allele Rht-D1b was also sometimes associated with reductions in early crop growth; no such negative effects were associated with the marker for Rht8c. When significant interactions between cropping system and genotype did occur it was because differences between lines were more exaggerated in the organic system than in the conventional system. Ppd-D1a was associated positively with plant numbers surviving the winter and early crop growth (GC, FR:R, DM, N, PAR, height), and was the most significant locus in a QTL analysis. We conclude that, within these environmental and system contexts, genes moderating development are likely to be more important in influencing early resource capture than using Rht8c as an alternative semi-dwarfing gene to Rht-D1b.

Influence of agricultural management, sward structure and food resources on grassland field use by birds in lowland England.

Agricultural management of grassland in lowland Britain has changed fundamentally in the last 50 years, resulting in spatial and structural uniformity within the pastoral landscape. The full extent to which these changes may have reduced the suitability of grassland as foraging habitat for birds is unknown. This study investigated the mechanisms by which these changes have impacted on birds and their food supplies. We quantified field use by birds in summer and winter in two grassland areas of lowland England (Devon and Buckinghamshire) over 3 years, relating bird occurrence to the management, sward structure and seed and invertebrate food resources of individual fields. Management intensity was defined in terms of annual nitrogen input. There was no consistent effect of management intensity on total seed head production, although those of grasses generally increased with inputs while forbs were rare throughout. Relationships between management intensity and abundance of soil and epigeal invertebrates were complex. Soil beetle larvae were consistently lower in abundance, and surface-active beetle larvae counts consistently higher, in intensively managed fields. Foliar invertebrates showed more consistent negatively relationships with management intensity. Most bird species occurred at low densities. There were consistent relationships across regions and years between the occurrence of birds and measures of field management. In winter, there was a tendency towards higher occupancy of intensively managed fields by species feeding on soil invertebrates. In summer, there were few such relationships, although many species avoided fields with tall swards. Use of fields by birds was generally not related to measures of seed or invertebrate food abundance. While granivorous species were perhaps too rare to detect a relationship, in insectivores the strong negative relationships (in summer) with sward height suggested that access to food may be the critical factor. While it appears that intensification of grassland management has been deleterious to the summer food resources of insectivorous birds that use insects living within the grass sward, intensification may have been beneficial to several species in winter through the enhancement of soil invertebrates. Synthesis and applications. We suggest that attempts to restore habitat quality for birds in grassland landscapes need to create a range of management intensities and sward structures at the field and farm scales. A greater understanding of methods to enhance prey accessibility, as well as abundance, for insectivorous birds is required.

Modelling habitat selection and distribution of the critically endangered Jerdon's courser Rhinoptilus bitorquatus in scrub jungle: an application of a new tracking method

1. Jerdon's courser Rhinoptilus bitorquatus is a nocturnally active cursorial bird that is only known to occur in a small area of scrub jungle in Andhra Pradesh, India, and is listed as critically endangered by the IUCN. Information on its habitat requirements is needed urgently to underpin conservation measures. We quantified the habitat features that correlated with the use of different areas of scrub jungle by Jerdon's coursers, and developed a model to map potentially suitable habitat over large areas from satellite imagery and facilitate the design of surveys of Jerdon's courser distribution. 2. We used 11 arrays of 5-m long tracking strips consisting of smoothed fine soil to detect the footprints of Jerdon's coursers, and measured tracking rates (tracking events per strip night). We counted the number of bushes and trees, and described other attributes of vegetation and substrate in a 10-m square plot centred on each strip. We obtained reflectance data from Landsat 7 satellite imagery for the pixel within which each strip lay. 3. We used logistic regression models to describe the relationship between tracking rate by Jerdon's coursers and characteristics of the habitat around the strips, using ground-based survey data and satellite imagery. 4. Jerdon's coursers were most likely to occur where the density of large (>2 m tall) bushes was in the range 300-700 ha(-1) and where the density of smaller bushes was less than 1000 ha(-1). This habitat was detectable using satellite imagery. 5. Synthesis and applications. The occurrence of Jerdon's courser is strongly correlated with the density of bushes and trees, and is in turn affected by grazing with domestic livestock, woodcutting and mechanical clearance of bushes to create pasture, orchards and farmland. It is likely that there is an optimal level of grazing and woodcutting that would maintain or create suitable conditions for the species. Knowledge of the species' distribution is incomplete and there is considerable pressure from human use of apparently suitable habitats. Hence, distribution mapping is a high conservation priority. A two-step procedure is proposed, involving the use of ground surveys of bush density to calibrate satellite image-based mapping of potential habitat. These maps could then be used to select priority areas for Jerdon's courser surveys. The use of tracking strips to study habitat selection and distribution has potential in studies of other scarce and secretive species.

The chromosomal assessment of salt tolerant substituted tritipyrum using genomic fluorescent in situ hybridiization (FISH)

Wheat, although moderately tolerant to salt, can not be cultivated in many areas. However, in the triticeae tribe, some of the wild wheat relatives are highly tolerant, e.g. Thinopyrum bessarabicum, which grows on the sea shore. Eight primary hexaploid tritipyrum lines, amphiploids between Triticum durum and Thinopyrum bessarabicum have been produced which can set seed in at least 250 mM NaCl. These tritipyrums (2n=6x=42, AABBEbEb) due to reasons such as brittle rachis, continuous production of tillers, late maturity, tall stature and meiotic instability will not fulfill the requirements of a successful commercial salt tolerant crop. To overcome such problems the substituted tritipyrum, in which selected Eb chromosomes are replaced by D genome chromosomes of 6x wheat, was produced from 6x tritipyrum x 6x wheat hybrids (F1: 2n=6x=42, AABBDEb) followed by selfing and backcrossing with 6x tritipyrum. The fertile plants among the above progenies were screened by the genomic fluorescent in situ hybridization technique to identify their Eb and D chromosome constitution. This study showed that producing tritiprum with variable numbers of Eb and D genome chromosomes is feasible and that FISH is a useful technique for determining the number of Eb chromosomes present.

Effects of reduced height (Rht) and photoperiod insensitivity (Ppd) alleles on yield of wheat in contrasting production systems

Near isogenic lines (NILs) varying for reduced height (Rht) and photoperiod insensitivity (Ppd-D1) alleles in a cv. Mercia background (rht (tall), Rht-B1b, Rht-D1b, Rht-B1c, Rht8c+Ppd-D1a, Rht-D1c, Rht12) were compared for interception of photosynthetically active radiation (PAR), radiation use efficiency (RUE), above-ground biomass (AGB), harvest index (HI), height, weed prevalence, lodging and grain yield, at one field site but within contrasting (‘organic’ v ‘conventional’) rotational and agronomic contexts, in each of three years. In the final year, further NILs (rht (tall), Rht-B1b, Rht-D1b, Rht-B1c, Rht-B1b+Rht-D1b, Rht-D1b+Rht-B1c) in Maris Huntsman and Maris Widgeon backgrounds were added together with 64 lines of a doubled haploid (DH) population [Savannah (Rht-D1b) × Renesansa (Rht-8c+Ppd-D1a)]. There were highly significant genotype × system interactions for grain yield, mostly because differences were greater in the conventional system than in the organic system. Quadratic fits of NIL grain yield against height were appropriate for both systems when all NILs and years were included. Extreme dwarfing was associated with reduced PAR, RUE, AGB, HI, and increased weed prevalence. Intermediate dwarfing was often associated with improved HI in the conventional system, but not in the organic system. Heights in excess of the optimum for yield were associated particularly with reduced HI and, in the conventional system, lodging. There was no statistical evidence that optimum height for grain yield varied with system although fits peaked at 85cm and 96cm in the conventional and organic systems, respectively. Amongst the DH lines, the marker for Ppd-D1a was associated with earlier flowering, and just in the conventional system also with reduced PAR, AGB and grain yield. The marker for Rht-D1b was associated with reduced height, and again just in the conventional system, with increased HI and grain yield. The marker for Rht8c reduced height, and in the conventional system only, increased HI. When using the System × DH line means as observations grain yield was associated with height and early vegetative growth in the organic system, but not in the conventional system. In the conventional system, PAR interception after anthesis correlated with yield. Savannah was the highest yielding line in the conventional system, producing significantly more grain than several lines that out yielded it in the organic system.

Effect of wheat dwarfing genes on nitrogen-use efficiency

Near isogenic lines (NILs) varying for alleles for reduced height (Rht) and photoperiod insensitivity (Ppd-D1a) in a cvar Mercia background (rht (tall), Rht-B1b, Rht-D1b, Rht-B1c, Rht8c+Ppd-D1a, Rht-D1c, Rht12) were compared at a field site in Berkshire, UK, but within different systems (‘organic’, O, in 2005/06, 2006/07 and 2007/08 growing seasons v. ‘conventional’, C, in 2005/06, 2006/07, 2007/08 and 2008/09). In 2007 and 2008, further NILs (rht (tall), Rht-B1b, Rht-D1b, Rht-B1c, Rht-B1b+Rht-D1b, Rht-D1b+Rht-B1c) in both Maris Huntsman and Maris Widgeon backgrounds were added. The contrasting systems allowed NILs to be tested in diverse rotational and agronomic, but commercially relevant, contexts, particularly with regard to the assumed temporal distribution of nitrogen availability, and competition from weeds. For grain, nitrogen-use efficiency (NUE; grain dry matter (DM) yield/available N; where available N=fertilizer N+soil mineral N), recovery of N in the grain (grain N yield/available N), N utilization efficiency to produce grain (NUtEg; grain DM yield/above-ground crop N yield), N harvest index (grain N yield/above-ground crop N yield) and dry matter harvest index (DMHI; grain DM yield/above-ground crop DM yield) all peaked at final crop heights of 800–950 mm. Maximum NUE occurred at greater crop heights in the organic system than in the conventional system, such that even adding just a semi-dwarfing allele (Rht-D1b) to the shortest background, Mercia, reduced NUE in the organic system. The mechanism of dwarfing (gibberellin sensitive or insensitive) made little difference to the relationship between NUE and its components with crop height. For above-ground biomass: dwarfing alleles had a greater effect on DM accumulation compared with N accumulation such that all dwarfing alleles could reduce nitrogen utilization efficiency (NUtE; crop DM yield/crop N yield). This was particularly evident at anthesis in the conventional system when there was no significant penalty for severe dwarfism for N accumulation, despite a 3-tonne (t)/ha reduction in biomass compared to the tallest lines. Differences between genotypes for recovery of N in the grain were thus mostly a function of net N uptake after anthesis rather than of remobilized N. This effect was compounded as dwarfing, except when coupled with Ppd-D1a, was associated with delayed anthesis. In the organic experiments there was greater reliance on N accumulated before anthesis, and genotype effects on NUE were confounded with effects on N accumulated by weeds, which was negatively associated with crop height. Optimum height for maximizing wheat NUE and its components, as manipulated by Rht alleles, thus depend on growing system, and crop utilization (i.e. biomass or grain production).

Semi-dwarfing (Rht-B1b) improves nitrogen-use efficiency in wheat, but not at economically optimal levels of nitrogen availability

A UK field experiment compared a complete factorial combination of three backgrounds (cvs Mercia, Maris Huntsman and Maris Widgeon), three alleles at the Rht-B1 locus as Near Isogenic Lines (NILs: rht-B1a (tall), Rht-B1b (semi-dwarf), Rht-B1c (severe dwarf)) and four nitrogen (N) fertilizer application rates (0, 100, 200 and 350 kg N/ha). Linear+exponential functions were fitted to grain yield (GY) and nitrogen-use efficiency (NUE; GY/available N) responses to N rate. Averaged over N rate and background Rht-B1b conferred significantly (P<0.05) greater GY, NUE, N uptake efficiency (NUpE; N in above ground crop / available N) and N utilization efficiency (NUtEg; GY / N in above ground crop) compared with rht-B1a and Rht-B1c. However the economically optimal N rate (Nopt) for N:grain price ratios of 3.5:1 to 10:1 were also greater for Rht-B1b, and because NUE, NUpE and NUtE all declined with N rate, Rht-Blb failed to increase NUE or its components at Nopt. The adoption of semi-dwarf lines in temperate and humid regions, and the greater N rates that such adoption justifies economically, greatly increases land-use efficiency, but not necessarily, NUE.

Reduced height alleles (Rht) and Hagberg falling number of wheat

Near isogenic lines varying for alleles for reduced height (Rht) and photoperiod insensitivity (Ppd-D1) in cv. Mercia (2005/6 to 2010/11; rht (tall), Rht-B1b, Rht-D1b, Rht-B1c, Rht8c+Ppd-D1a, Rht-D1c, Rht12) and cvs Maris Huntsman and Maris Widgeon (2007/8 to 2010/11; rht (tall), Rht-B1b, Rht-D1b, Rht-B1c, Rht-B1b+Rht-D1b, Rht-D1b+Rht-B1c) were compared at one field site, but within different systems (‘organic’, O, 2005/6 to 2007/8 v ‘intensive’, I, 2005/6 to 2010/11). Further experiments at the site (2006/7 to 2008/9) compared 64 lines of a doubled haploid (DH) population [Savannah (Rht-D1b) × Renesansa (Rht-8c+Ppd-D1a)]. Gibberellin (GA) insensitive dwarfing alleles (Rht-B1b; Rht-B1c; Rht-D1b; Rht-D1c) could reduce α-amylase activity and/or increase Hagberg falling number (HFN) but effects depended greatly on system, background and season. Only Rht-B1c increased grain dormancy despite producing plants taller than Rht-D1c. The GA-sensitive Rht8c+Ppd-D1a in Mercia was associated with reduced HFN but analysis of the DH population suggested this was more closely linked with Ppd-D1a, rather than Rht8c. The severe GA-sensitive dwarfing allele Rht12 was associated with reduced HFN. Instability in HFN over season tended to increase with degree of dwarfing. There was a negative association between mean grain weight and HFN that was in addition to effects of Rht and Ppd-D1 allele.

Under what conditions does embedded convection enhance orographic precipitation?

Idealised convection-permitting simulations are used to quantify the impact of embedded convection on the precipitation generated by moist flow over midlatitude mountain ridges. A broad range of mountain dimensions and moist stabilities are considered to encompass a spectrum of physically plausible flows. The simulations reveal that convection only enhances orographic precipitation in cap clouds that are otherwise unable to efficiently convert cloud condensate into precipitate. For tall and wide mountains (e.g. the Washington Cascades or the southern Andes), precipitate forms efficiently through vapour deposition and collection, even in the absence of embedded convection. When embedded convection develops in such clouds, it produces competing effects (enhanced condensation in updraughts and enhanced evaporation through turbulent mixing and compensating subsidence) that cancel to yield little net change in precipitation. By contrast, convection strongly enhances precipitation over short and narrow mountains (e.g. the UK Pennines or the Oregon Coastal Range) where precipitation formation is otherwise highly inefficient. Although cancellation between increased condensation and evaporation still occurs, the enhanced precipitation formation within the convective updraughts leads to a net increase in precipitation efficiency. The simulations are physically interpreted through non-dimensional diagnostics and relevant time-scales that govern advective, microphysical, and convective processes.

Gibberellin-responsive and -insensitive dwarfing alleles on wheat performance in contrasting tillage systems

Near-isogenic lines (NILs) of winter wheat varying for alleles for reduced height (Rht), gibberellin (GA) response and photoperiod insensitivity (Ppd-D1a) in cv. Mercia background (rht (tall), Rht-B1b, Rht-D1b, Rht-B1c, Rht8c+Ppd-D1a, Rht-D1c, Rht12) and cv. Maris Widgeon (rht (tall), Rht-D1b, Rht-B1c) backgrounds were compared to investigate main effects and interactions with tillage (plough-based, minimum-, and zero-tillage) over two years. Both minimum- and zero- tillage were associated with reduced grain yields allied to reduced harvest index, biomass accumulation, interception of photosynthetically active radiation (PAR), and plant populations. Grain yields were optimized at mature crop heights of around 740mm because this provided the best compromise between harvest index which declined with height, and above ground biomass which increased with height. Improving biomass with height was due to improvements in both PAR interception and radiation-use efficiency. Optimum height for grain yield was unaffected by tillage system or GA-sensitivity. After accounting for effects of height, GA insensitivity was associated with increased grain yields due to increased grains per spike, which was more than enough to compensate for poorer plant establishment and lower mean grain weights compared to the GA-sensitive lines. Although better establishment was possible with GA-sensitive lines, there was no evidence that this effect interacted with tillage method. We find, therefore, little evidence to question the current adoption of wheats with reduced sensitivity to GA in the UK, even as tillage intensity lessens.

Isolation and characterization of an AINTEGUMENTA-like gene in different coconut (Cocos nucifera L.) varieties from Sri Lanka

Development of an efficient tissue culture protocol in coconut is hampered by numerous technical constraints. Thus a greater understanding of the fundamental aspects of embryogenesis is essential. The role of AINTEGUMENTA-like genes in embryogenesis has been elucidated not only in model plants but also in economically important crops. A coconut gene, CnANT, that encodes two APETALA2 (AP2) domains and a conserved linker region similar to those of the BABY BOOM transcription factor was cloned, characterized, and its tissue specific expression was examined. The full-length cDNA of 1,780 bp contains a 1,425-bp open reading frame that encodes a putative peptide of 474 amino acids. The genomic DNA sequence includes 2,317 bp and consists of nine exons interrupted by eight introns. The exon/intron organization of CnANT is similar to that of homologous genes in other plant species. Analysis of differential tissue expression by real-time polymerase chain reaction indicated that CnANT is expressed more highly in in vitro grown tissues than in other vegetative tissues. Sequence comparison of the genomic sequence of CnANT in different coconut varieties revealed one single nucleotide polymorphism and one indel in the first exon and first intron, respectively, which differentiate the Tall group of trees from Dwarfs. The indel sequence, which can be considered a simple sequence repeats marker, was successfully used to distinguish the Tall and Dwarf groups as well as to develop a marker system, which may be of value in the identification of parental varieties that are used in coconut breeding programs in Sri Lanka.

An assessment of a three-beam Doppler lidar wind profiling method for use in urban areas

Currently there are few observations of the urban wind field at heights other than rooftop level. Remote sensing instruments such as Doppler lidars provide wind speed data at many heights, which would be useful in determining wind loadings of tall buildings, and predicting local air quality. Studies comparing remote sensing with traditional anemometers carried out in flat, homogeneous terrain often use scan patterns which take several minutes. In an urban context the flow changes quickly in space and time, so faster scans are required to ensure little change in the flow over the scan period. We compare 3993 h of wind speed data collected using a three-beam Doppler lidar wind profiling method with data from a sonic anemometer (190 m). Both instruments are located in central London, UK; a highly built-up area. Based on wind profile measurements every 2 min, the uncertainty in the hourly mean wind speed due to the sampling frequency is 0.05–0.11 m s−1. The lidar tended to overestimate the wind speed by ≈0.5 m s−1 for wind speeds below 20 m s−1. Accuracy may be improved by increasing the scanning frequency of the lidar. This method is considered suitable for use in urban areas.

Observations of wind speed profiles over Greater London, UK, using a Doppler lidar

To calculate the potential wind loading on a tall building in an urban area, an accurate representation of the wind speed profile is required. However, due to a lack of observations, wind engineers typically estimate the characteristics of the urban boundary layer by translating the measurements from a nearby reference rural site. This study presents wind speed profile data obtained from a Doppler lidar in central London, UK, during an 8 month observation period. Used in conjunction with wind speed data measured at a nearby airport, the data have been used to assess the accuracy of the predictions made by the wind engineering tools currently available. When applied to multiple changes in surface roughness identified from morphological parameters, the non-equilibrium wind speed profile model developed by Deaves (1981) provides a good representation of the urban wind speed profile. For heights below 500 m, the predicted wind speed remains within the 95% confidence interval of the measured data. However, when the surface roughness is estimated using land use as a proxy, the model tends to overestimate the wind speed, particularly for very high wind speed periods. These results highlight the importance of a detailed assessment of the nature of the surface when estimating the wind speed above an urban surface.

Effect of Rht alleles on the tolerance of wheat grain set to high temperature and drought stress during booting and anthesis

Factorial pot experiments were conducted to compare the responses of GA-sensitive and GA-insensitive reduced height (Rht) alleles in wheat for susceptibility to heat and drought stress during booting and anthesis. Grain set (grains/spikelet) of near isogenic lines (NILs) was assessed following three day transfers to controlled environments imposing day temperatures (t) from 20 to 40°C. Transfers were during booting and/or anthesis and pots maintained at field capacity (FC) or had water withheld. Logistic responses (y = c/1+e-b(t -m)) described declining grain set with increasing t, and t5 was that fitted to give a 5% reduction in grain set. Averaged over NIL, t5 for anthesis at FC was 31.7±0.47°C (S.E.M, 26 d.f.). Drought at anthesis reduced t5 by <2°C. Maintaining FC at booting conferred considerable resistance to high temperatures (t5=33.9°C) but booting was particularly heat susceptible without water (t5 =26.5°C). In one background (cv. Mercia), for NILs varying at the Rht-D1 locus, there was progressive reduction in t5 with dwarfing and reduced gibberellic acid (GA) sensitivity (Rht-D1a, tall, 32.7±0.72; Rht-D1b, semi-dwarf, 29.5±0.85; Rht-D1c, severe dwarf, 24.2±0.72). This trend was not evident for the Rht-B1 locus, or for Rht-D1b in an alternative background (Maris Widgeon). The GA-sensitive severe dwarf Rht12 was more heat tolerant (t5=29.4±0.72) than the similarly statured GA-insensitive Rht-D1c. The GA-sensitive, semi-dwarfing Rht8 conferred greater drought tolerance in one experiment. Despite the effects of Rht-D1 alleles in Mercia on stress tolerance, the inconsistency of the effects over background and locus led to the conclusion that semi-dwarfing with GA-insensitivity did not necessarily increase sensitivity to stress at booting and flowering. In comparison to effects of semi-dwarfing alleles, responses to heat stress are much more dramatically affected by water availability and the precise growth stage at which the stress is experienced by the plants.

Ectomycorrhizal symbiosis can enhance plant nutrition through improved access to discrete organic nutrient patches of high resource quality

It is known that roots can respond to patches of fertility; however, root proliferation is often too slow to exploit resources fully, and organic nutrient patches may be broken down and leached, immobilized or chemically fixed before they are invaded by the root system. The ability of fungal hyphae to exploit resource patches is far greater than that of roots due to their innate physiological and morphological plasticity, which allows comprehensive exploration and rapid colonization of resource patches in soils. The fungal symbionts of ectomycorrhizal plants excrete significant quantities of enzymes such as chitinases, phosphatases and proteases. These might allow the organic residue to be tapped directly for nutrients such as N and P. Pot experiments conducted with nutrient-stressed ectomycorrhizal and control willow plants showed that when high quality organic nutrient patches were added, they were colonized rapidly by the ectomycorrhizal mycelium. These established willows (0.5 m tall) were colonized by Hebeloma syrjense P. Karst. for 1 year prior to nutrient patch addition. Within days after patch addition, colour changes in the leaves of the mycorrhizal plants (reflecting improved nutrition) were apparent, and after I month the concentration of N and P in the foliage of mycorrhizal plants was significantly greater than that in non-mycorrhizal plants subject to the same nutrient addition. It seems likely that the mycorrhizal plants were able to compete effectively with the wider soil microbiota and tap directly into the high quality organic resource patch via their extra-radical mycelium. We hypothesize that ectomycorrhizal plants may reclaim some of the N and P invested in seed production by direct recycling from failed seeds in the soil. The rapid exploitation of similar discrete, transient, high-quality nutrient patches may have led to underestimations when determining the nutritional benefits of ectomycorrhizal colonization.