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Vertical transmission of symbiotic Epichloë endophytes from host grasses into progeny seed is the primary mechanism by which the next generation of plants is colonized. This process is often imperfect, resulting in endophyte-free seedlings which may have poor ecological fitness if the endophyte confers protective benefits to its host. In this study, we investigated the influence of host genetics and environment on the vertical transmission of Epichloë festucae var. lolii strain AR37 in the temperate forage grass Lolium perenne . The efficiency of AR37 transmission into the seed of over 500 plant genotypes from five genetically diverse breeding populations was determined. In Populations I-III, which had undergone previous selection for high seed infection by AR37, mean transmission was 88, 93, and 92%, respectively. However, in Populations IV and V, which had not undergone previous selection, mean transmission was 69 and 70%, respectively. The transmission values, together with single-nucleotide polymorphism data obtained using genotyping-by-sequencing for each host, was used to develop a genomic prediction model for AR37 seed transmission. The predictive ability of the model was estimated at r = 0.54. While host genotype contributed greatly to differences in AR37 seed transmission, undefined environmental variables also contributed significantly to seed transmission across different years and geographic locations. There was evidence for a small host genotype-by-environment effect; however this was less pronounced than genotype or environment alone. Analysis of endophyte infection levels in parent plants within Populations I and IV revealed a loss of endophyte infection over time in Population IV only. This population also had lower average tiller infection frequencies than Population I, suggesting that AR37 failed to colonize all the daughter tillers and therefore seeds. However, we also observed that infection of seed by AR37 may fail during or after initiation of floral development from plants where all tillers remained endophyte-infected over time. While the effects of environment and host genotype on fungal endophyte transmission have been evaluated previously, this is the first study that quantifies the relative impacts of host genetics and environment on endophyte vertical transmission.

Key Message: Genomic prediction models for multi-year dry matter yield, via genotyping-by-sequencing in a composite training set, demonstrate potential for genetic gain improvement through within-half sibling family selection. Perennial ryegrass (Lolium perenne L.) is a key source of nutrition for ruminant livestock in temperate environments worldwide. Higher seasonal and annual yield of herbage dry matter (DMY) is a principal breeding objective but the historical realised rate of genetic gain for DMY is modest. Genomic selection was investigated as a tool to enhance the rate of genetic gain. Genotyping-by-sequencing (GBS) was undertaken in a multi-population (MP) training set of five populations, phenotyped as half-sibling (HS) families in five environments over 2 years for mean herbage accumulation (HA), a measure of DMY potential. GBS using the ApeKI enzyme yielded 1.02 million single-nucleotide polymorphism (SNP) markers from a training set of n = 517. MP-based genomic prediction models for HA were effective in all five populations, cross-validation-predictive ability (PA) ranging from 0.07 to 0.43, by trait and target population, and 0.40-0.52 for days-to-heading. Best linear unbiased predictor (BLUP)-based prediction methods, including GBLUP with either a standard or a recently developed (KGD) relatedness estimation, were marginally superior or equal to ridge regression and random forest computational approaches. PA was principally an outcome of SNP modelling genetic relationships between training and validation sets, which may limit application for long-term genomic selection, due to PA decay. However, simulation using data from the training experiment indicated a twofold increase in genetic gain for HA, when applying a prediction model with moderate PA in a single selection cycle, by combining among-HS family selection, based on phenotype, with within-HS family selection using genomic prediction.

This study tested the hypotheses that: (i) genetic variation in Rubisco turnover may exist in perennial ryegrass (Lolium perenne L.); (ii) such variation might affect nitrogen use efficiency and plant yield; and (iii) genetic control of Rubisco turnover might be amenable to identification by quantitative trait loci (QTL) mapping. A set of 135 full-sib F1 perennial ryegrass plants derived from a pair cross between genotypes from the cultivars 'Grasslands Impact' and 'Grasslands Samson' was studied to test these hypotheses. Leaf Rubisco concentration at different leaf ages was measured and modelled as a log-normal curve described by three mathematical parameters: D (peak Rubisco concentration), G (time of D), and F (curve standard deviation). Herbage dry matter (DM) yield and morphological traits (tiller weight (TW), tiller number (TN), leaf lamina length (LL), and an index of competitive ability (PI)) were also measured. The progeny exhibited continuous variation for all traits. Simple correlation and principal component analyses indicated that plant productivity was associated with peak Rubisco concentration and not Rubisco turnover. Lower DM was associated with higher leaf Rubisco concentration indicating that Rubisco turnover effects on plant productivity may relate to energy cost of Rubisco synthesis rather than photosynthetic capacity. QTL detection by a multiple QTL model identified seven significant QTL for Rubisco turnover and nine QTL for DM and morphological traits. An indication of the genetic interdependence of DM and the measures of Rubisco turnover was the support interval overlap involving QTL for D and QTL for TN on linkage group 5 in a cluster involving QTL for DM and PI. In this region, alleles associated with increased TN, DM, and PI were associated with decreased D, indicating that this region may regulate Rubisco concentration and plant productivity via increased tillering. A second cluster involving QTL for LL, TN, PI and DM was found on linkage group 2. The two clusters represent marker-trait associations that might be useful for marker-assisted plant breeding applications. In silico comparative analysis indicated conservation of the genetic loci controlling Rubisco concentration in perennial ryegrass and rice.

Endophytic fungi in pasture grasses produce alkaloids which affect invertebrate and vertebrate herbivores. While the competence to produce an alkaloid is a property of the fungus, the host plant may moderate fungal activity. Host genetic influence on endophyte activity was studied in perennial ryegrass (Lolium perenne L.) infected with a common strain of Neotyphodium lolii (Latch, Christensen & Samuels) Glenn, Bacon & Hanlin. Progeny seedling families of a partial diallel cross and their 12 parent clones were compared in a glasshouse experiment. Peramine and ergovaline concentrations were determined by high pressure liquid chromatography (HPLC), and intensity of endophyte infection was determined by enzyme-linked immunosorbent assay (ELISA). Concentrations of peramine and ergovaline and the amount of endophyte mycelium in plants varied between families, consistently across two glasshouse cells and (for the HPLC data) two harvests. There was no indication of any maternal effects. Host genetic control was evident in significant general combining ability effects and smaller specific combining ability effects. Parent-progeny correlation coefficients were high, and narrow-sense heritability was estimated as 0.70, 0.72, and 0.58 respectively for ergovaline, peramine, and ELISA. Further analysis indicated little interaction between loci, and no directional dominance. The three traits were correlated, indicating that 41 and 65% of the genetically controlled variation in ergovaline and peramine concentrations, respectively, was a function of mycelial mass. However, there were departures from these relationships. Host plant selection may enable development of pastures with controlled low levels of toxic but ecologically beneficial endophyte metabolites.

The concept of the morphogenetic field has been used extensively in developmental biology. However, little is known about the mechanisms that partition these broad areas of tissue into the smaller areas which actually form the corresponding structures, and the remaining tissue. In the Mexican axolotl, the heart field forms as the anterior lateral plate mesoderm migrates over the underlying pharyngeal endoderm between stages 14 and 28. We have previously shown that both the mid-ventral and lateral walls of the pharyngeal cavity have considerable inductive capacity at stage 14. If this inductive capability, and the competence of the mesoderm to respond, is retained between stages 14 and 28, a much broader area of mesoderm would be induced than actually participates in heart development. In this paper, we use explant cultures to establish that pharyngeal endoderm retains its inductive activity, and that both pre-cardiac mesoderm and lateral plate mesoderm caudal to the pharyngeal cavity remain competent to respond to the induction throughout this period. We also map the specified region of the antero-lateral mesoderm between stages 14 and 28 by placing carefully measured areas of mesoderm in culture without inductive endoderm. We found that the region capable of initiating a spontaneous beat approximately doubles in size during this period. Since the specified region is larger than the actual heart primordium, some mechanism must exist to partition "induced" mesoderm into heart-forming and non-heart-forming areas.(ABSTRACT TRUNCATED AT 250 WORDS)

The chick heart is formed from bilateral patches of presumptive cardiac mesoderm cells which migrate over the endoderm and fuse in the midline. We have tested the possibility that this migration is controlled, at least in part, by a chemotactic substance exuded by the anterior end of the endoderm. We have used chick/quail combinations to follow naturally marked cells during the course of their migration. Chimaeric embryos were formed by fusing together parts of chick and quail embryos of stage 5-6. Each embryo possessed two pairs of precardiac regions, the quail pair lying immediately anterior to that of the chick. These chimaeras were then explanted in embryo culture. In the event of chemotaxis, cells from the posterior end of the quail precardiac mesoderm might be expected to invade the chick area. Samples of explants and chimaeras were examined at intervals from 2 to 24 h, but in no case were cells found to have changed their direction of migration as a result of the proximity of anterior endoderm. It is concluded that this work does not provide evidence for a chemotactic attraction by the anterior end of the endoderm.