First Report of Powdery Mildew Caused by Golovinomyces ambrosiae on Xanthium orientale in Korea.

Xanthium orientale L. (syn. Xanthium canadense Mill., Asteraceae), known as cocklebur, is an annual weed native to North America, which is now a neophyte distributed throughout the world. This plant was accidentally introduced to Korea in the late 1970s ( So et al. 2008) and is considered a problematic exotic weed in orchards, for which many herbicides are ineffective (Kim et al. 2020). In September 2018, powdery mildew was observed on X. orientale in Jeju, Korea. The disease incidence ranged from 40 to 60%. Voucher specimens were deposited in the Korea University Herbarium (Accession No. KUS-F30795) and Kunsan National University Herbarium (KSNUH1988). Symptoms appeared as round to irregular white patches with abundant hyphal growth on the leaf surface. Hyphal appressoria were nipple-shaped, and 3 to 6 µm diam. Conidiophores (n = 30) were 145 to 206 × 9 to 11.6 µm and produced 2 to 5 immature conidia in chains with a sinuate outline. Foot-cells of the conidiophores were straight, cylindrical, and 43 to 100.9 µm long. Conidia (n = 30) were ellipsoid-ovoid, doliiform to somewhat limoniform, 25.2 to 31.8 × 13.6 to 16.8 µm (l/w 1.6 to 2.1), and devoid of distinct fibrosin bodies. The morphological characteristics corresponded to those of Golovinomyces ambrosiae (Schwein.) U. Braun & R.T.A. Cook (Braun and Cook 2012, under Golovinomyces spadiceus (Beck. & M.A. Curtis) U. Braun; Qiu et al. 2020). To confirm the identity of the causal fungus, the internal transcribed spacer (ITS), large subunit (LSU) (Bradshaw and Tobin 2020), the intergenic spacer (IGS) of rDNA, and the glyceraldehyde-3-phosphate dehydrogenase (GAPDH) gene (Bradshaw et al. 2022) were amplified for a herbarium specimen (KUS-F30795). A BLASTn search of these sequences revealed 100% identity with reference sequences of G. ambrosiae on diverse Asteraceae plants (AB077644 for ITS, AB077643 for LSU, ON361171 for IGS, and ON075648 for GAPDH). However, there was a single nucleotide difference on both the IGS and GAPDH sequences when compared to the closely related species Golovinomyces latisporus. The sequences were deposited in GenBank (Accession No. OQ165157 (ITS), OQ165164 (LSU), OR050524 (IGS), and OR086076 (GAPDH)). Phylogenetic analyses of ITS, LSU, IGS, and GAPDH sequences revealed the Korean sample formed a well-supported group with other G. ambrosiae sequences, confirming its identity. A pathogenicity test was performed through inoculation by gently pressing diseased leaves onto the leaves of five healthy plants. Five non-inoculated plants served as controls. All plants were maintained in a greenhouse at 25±2°C. Powdery mildew colonies developed on the inoculated plants after ten days, whereas the control plants remained symptomless. The fungus present on the inoculated leaves was morphologically identical to that observed on the initially diseased leaves, fulfilling Koch's postulates. Powdery mildew on X. orientale has previously been reported as Golovinomyces cichoracearum (≡ Erysiphe cichoracearum) sensu lato in the USA, G. ambrosiae (= G. spadiceus) throughout all continents, and Podosphaera fusca sensu lato (now P. xanthii) in Korea (Braun and Cook 2012; Farr and Rossman 2023). To date, powdery mildew in Korea has been reported only on Xanthium strumarium as G. cichoracearum s. lat. and Podosphaera xanthii (KSPP 2022). To our knowledge, this is the first report of powdery mildew caused by G. ambrosiae on X. orientale in Korea.

Inconsistent intraspecific pattern in leaf life span along nitrogen-supply gradient.

PREMISE OF THE STUDY: Leaf life span (LLS) has long been hypothesized to plastically increase with decreasing nitrogen (N) supply from soil to maximize N retention, carbon assimilation, and fitness; however, accumulating evidence shows no consistent trend. The apparent inconsistencies are explained by a recent model that assumes LLS has a hump-shaped quadratic response to the N-supply gradient. The available evidence mostly originates from comparisons of LLS at only two levels of N availability, and the hypothesis remains unanswered. METHODS: We investigated LLS of two asteraceous forbs (Adenocaulon himalaicum and Xanthium canadense) experimentally grown at eight levels of N supply, which covered a range of N supply in their natural habitats. We additionally conducted a literature search to retrieve studies reporting LLS response along an N-supply gradient. KEY RESULTS: The LLS of neither species showed a hump-shaped response along the N-supply gradient. Past studies examining the LLS of an aquatic forb and terrestrial shrubs and trees along the N-supply gradient (more than four levels of N supply) also refuted the hypothesis. CONCLUSIONS: The LLS of a single species exhibited neither an increase nor a hump-shaped response to decreased N supply in a variety of life forms. Comparisons at only a few N levels are misleading with regard to LLS response to N supply.

Leaf dynamics in growth and reproduction of Xanthium canadense as influenced by stand density.

BACKGROUND AND AIMS: Leaf longevity is controlled by the light gradient in the canopy and also by the nitrogen (N) sink strength in the plant. Stand density may influence leaf dynamics through its effects on light gradient and on plant growth and reproduction. This study tests the hypothesis that the control by the light gradient is manifested more in the vegetative period, whereas the opposite is true when the plant becomes reproductive and develops a strong N sink. METHODS: Stands of Xanthium canadense were established at two densities. Emergence, growth and death of every leaf on the main stem and branches, and plant growth and N uptake were determined from germination to full senescence. Mean residence time and dry mass productivity were calculated per leaf number, leaf area, leaf mass and leaf N (collectively termed 'leaf variables') in order to analyse leaf dynamics and its effect on plant growth. KEY RESULTS: Branching and reproductive activities were higher at low than at high density. Overall there was no significant difference in mean residence time of leaf variables between the two stands. However, early leaf cohorts on the main stem had a longer retention time at low density, whereas later cohorts had a longer retention time at high density. Branch leaves emerged earlier and tended to live longer at low than at high density. Leaf efficiencies, defined as carbon export per unit investment of leaf variables, were higher at low density in all leaf variables except for leaf number. CONCLUSIONS: In the vegetative phase of plant growth, the light gradient strongly controls leaf longevity, whereas later the effects of branching and reproductive activities become stronger and over-rule the effect of light environment. As leaf N supports photosynthesis and also works as an N source for plant development, N use is pivotal in linking leaf dynamics with plant growth and reproduction.

Stem extension and mechanical stability of Xanthium canadense grown in an open or in a dense stand.

BACKGROUND AND AIMS: Plants in open, uncrowded habitats typically have relatively short stems with many branches, whereas plants in crowded habitats grow taller and more slender at the expense of mechanical stability. There seems to be a trade-off between height growth and mechanical stability, and this study addresses how stand density influences stem extension and consequently plant safety margins against mechanical failure. METHODS: Xanthium canadense plants were grown either solitarily (S-plants) or in a dense stand (D-plants) until flowering. Internode dimensions and mechanical properties were measured at the metamer level, and the critical buckling height beyond which the plant elastically buckles under its own weight and the maximum lateral wind force the plant can withstand were calculated. KEY RESULTS: Internodes were longer in D- than S-plants, but basal diameter did not differ significantly. Relative growth rates of internode length and diameter were negatively correlated to the volumetric solid fraction of the internode. Internode dry mass density was higher in S- than D-plants. Young's modulus of elasticity and the breaking stress were higher in lower metamers, and in D- than in S-plants. Within a stand, however, both moduli were positively related to dry mass density. The buckling safety factor, a ratio of critical buckling height to actual height, was higher in S- than in D-plants. D-plants were found to be approaching the limiting value 1. Lateral wind force resistance was higher in S- than in D-plants, and increased with growth in S-plants. CONCLUSIONS: Critical buckling height increased with height growth due mainly to an increase in stem stiffness and diameter and a reduction in crown/stem mass ratio. Lateral wind force resistance was enhanced due to increased tissue strength and diameter. The increase in tissue stiffness and strength with height growth plays a crucial role in maintaining a safety margin against mechanical failure in herbaceous species that lack the capacity for secondary growth.

Growth schedule of Xanthium canadense: Does it optimize the timing of reproduction?

The effects of nutrition on the timing of reproductive initiation of a short-day annual plant Xanthium canadense (cocklebur) were examined with the following hypotheses in mind: If the plant always follows an optimal growth schedule, low-nutrient plants will initiate reproductive growth earlier than high-nutrient plants. On the other hand, if the plant flowers in response to photoperiodic stimuli, both plants will initiate reproductive growth on the same day. The sand-culture experiment showed that high-nutrient plants flowered earlier than the low-nutrient plants, leading to rejection of the first hypothesis. The predicted optimal flowering time is 2 days later than the actual flowering time in high-nutrient plants and 10 days earlier in low-nutrient plants. These deviations from the optimal times reduced the reproductive yield by 0.1% and 2.3%, respectively. The ratio of the final reproductive yield to the vegetative mass at flower initiation was 1.10 in high-nutrient plants and 0.63 in low-nutrient plants. Since the expected ratio for the optimal growth schedule is 1.0, high-nutrient plants followed the opitmal growth schedule more closely than the low-nutrient plants. Cocklebur is a fast-growing annual which is common in relatively nutrient-rich environments. This study suggests that cocklebur adapts itself to such environments through its photoperiodic response.

Timing of seed germination and the reproductive effort in Xanthium canadense.

The effect of different dates of germination on the timing of flowering and the final reproductive yield was examined in a short-day annual plant Xanthium canadense (cocklebur). Delays in germination of 30 and 60 days deferred flower initiation by 2 and 9 days, respectively. Although plants that germinated later were smaller because of the shorter growing period, the reproductive yields did not show as much reduction as the vegetative biomass. The reproductive effort (RE, defined as the ratio of final reproductive yield to the vegetative biomass at the end of the growing season) increased 1.5 and 2.5 times with delays in germination of 30 and 60 days, respectively. A simple model of plant growth was used to analyse the factors involved in the control of RE, which depends only on the dry mass productivity and its partitioning in the reproductive phase, and is independent of the productivity and partitioning in the vegetative phase. Since relative allocation of dry mass to the reproductive part in the reproductive phase was similar for plants with different germination dates, the different REs could be ascribed mainly to differences in productivity of the vegetative parts in the reproductive period. The dependence of RE on plant size is discussed.

Effects of shift in flowering time on the reproductive output of Xanthium canadense in a seasonal environment.

We studied the effects of a change in flowering date on the reproductive output of a short-day annual plant, Xanthium canadense. The flowering date was changed by photoperiodic manipulation to 1 month earlier or later than the natural flowering date. Plants with the natural flowering date attained the highest reproductive output. For those flowering 1 month earlier or later, the reproductive output was decreased by 42% or 23%, respectively. The reproductive output was analyzed as the product of the biomass production during the reproductive period and its allocation to the reproductive organs. Although delay in flowering increased biomass production, it decreased its fractional allocation to the reproductive organs. The highest reproductive output in the natural flowering plants resulted from a compromise between these two effects of flowering. Plants flowering earlier had higher translocation rates to the reproductive organs and accelerated plant senescence. Later flowering caused a reduction in biomass translocation to the reproductive organs and thus extended the reproductive period. These experimental results are discussed in relation to the cost of reproduction and the optimal time for flowering that maximizes the final reproductive output. It is suggested that the natural flowering time maximized the reproductive output while minimizing the cost of reproduction.

Leaf nitrogen distribution in relation to leaf age and photon flux density in dominant and subordinate plants in dense stands of a dicotyledonous herb.

We studied the effects of photon flux density (PFD) and leaf position, a measure of developmental age, on the distribution of nitrogen content per unit leaf area (N area) in plants of different heights, in dense stands grown at two nitrogen availabilities and in solitary plants of the erect dicotyledonous herb Xanthium canadense. Taller more dominant plants received higher PFD levels and experienced a larger difference in relative PFD between their youngest and oldest leaves than shorter subordinate plants in the stands. Differences in PFD between leaves of solitary plants were assumed to be minimal and differences in leaf traits, found for these plants, could thus be mainly attributed to an effect of leaf position. In the solitary plants, N area decreased with leaf position while in the plants from the stands it decreased with decreasing relative PFD, indicating both factors to be important in determining the distribution of N area. Due to the effect of leaf position on N area, leaves of subordinate plants had a higher N area than older leaves of dominant plants which were at the same height or slightly higher in the canopy. Consequently, the N area distribution patterns of individual plants plotted as a function of relative PFD were steeper, and probably closer to the optimal distribution which maximizes photosynthesis, than the average distribution in the stand. Leaves of subordinate plants had a lower mass per unit area (LMA) than those of dominant plants. In the dominant plants, LMA decreased with decreasing relative PFD (and with leaf position) while in the subordinate plants it increased. This surprising result for the subordinate plants can be explained by the fact that, during the course of a growing season, these plants became increasingly shaded and newer leaves were thus formed at progressively lower light availability. This indicates that LMA was strongly determined by the relative PFD at leaf formation and to a lesser extent by the current PFD. Leaf N content per unit mass (N mass) was strongly determined by leaf position independent of relative PFD. This indicates that N mass is strongly ontogenetically related to the leaf-aging process while changes in N area, in response to PFD, were regulated through changes in LMA.