Application of the equilibrium concept to the development of agaric fruit-bodies, with special reference to their straight downward growth in light from below

Equilibrium, a concept of dynamics, is found to be applicable to the phototropic and gravitropic growth in agaric fruit-bodies. The fruit-bodies exposed to light from below grow straight downward without bending upward, and those exposed to light from obliquely below grow first downward and then upward by negative gravitropism. The fruit-bodies exposed to light from above grow upward. Fruit-bodies growing straight downward or upward do not change the direction of growth; they are in ‘equilibria’. The straight downward growth can be regarded as an ‘unstable equilibrium’ having a higher potential, and the straight upward growth as a ‘stable equilibrium’ having a lower potential. The change in the direction of growth can be explained by the change in the potential; the upward bending in fruit-bodies that have grown obliquely downward can be regarded as a ‘transition’ from the unstable equilibrium to the stable one.     

Responses of agaric fruit-bodies to light and gravity: growth straight downward in response to light from below

Fruit-bodies of Agaricales are known to show positive phototropism during the early stage of development, but negative gravitropism at the later stage after the onset of basidiospore formation. However, when exposed to light from below, the fruit-bodies ofTephrocybe tesquorum andCoprinus spp. grew downward through all stages of development, even after the onset of basidiospore formation. Primordium formation, fruit-body development and basidiospore formation were not disturbed under such conditions. In these downward-growing fruit-bodies, gills stood straight upward. InT. tesquorum, caps often became swollen and stipes sometimes became twisted anticlockwise, contrary to those in light from above, while such behaviours were not observed inCoprinus spp.     

Responses of lignicolous-agaric fruit-bodies to light and gravity: A study to overview the fruit-body development in hymenomycetes

Our previous study revealed that, when exposed to light from below, fruit-bodies of humus-borne agarics grow straight downward both in the ‘Coprinus Type’ and ’Non-Coprinus Type’ species and that, in the latter, gills tilt by gravitropism, caps swell and wave, and stipes twist. The present study revealed that fruit-bodies of some lignicolous agarics also grew straight downward. Among themFlammulina velutipes andHypsizygus marmoreus fruit-bodies showed almost the same behaviour as the ‘Non-Coprinus Type’, butPleurotus ostreatus (assumedPolyporus Type) fruit-bodies did not show the gill tilting and cap waving. Rather, it redifferentiated to produce new gills or new fruit-bodies, possibly by gravimorphogenesis, or formed spiral stipes. Based on these results, graviresponses in hymenomycetes are overviewed.     

Growth dynamics of Potamogeton pectinatus L. in Lake Burullus,Egypt: a modelling approach

In this study, we used a macrophyte model to describe the growth production and the interaction between above‐ and below‐ground organs of Potamogeton pectinatus in Lake Burullus, Egypt. Above‐ and below‐ground biomass of P. pectinatus was sampled on a monthly basis from April to December 2011 at three sites of Lake Burullus. Shoots started to grow in April, reached the maximum biomass in September and then rapidly decreased in October when they moved into the senescence stage. Tubers biomass reduced in August due to the upward translocation to shoots, but sharply increased to the maximum in October by downward translocation from shoots and roots. Potamogeton pectinatus allocated approximately 82.3% of its total biomass to shoots, 15.5% to tubers and 2.2% to roots.     

Linking above-ground and below-ground effects in autotrophic microcosms: effects of shading and defoliation on plant and soil properties

Although factors affecting plant growth and plant carbon/nutrient balance – e.g., light availability and defoliation by herbivores – may also propagate changes in below‐ground food webs, few studies have aimed at linking the above‐ground and below‐ground effects. We established a 29‐week laboratory experiment (～one growing season) using autotrophic microcosms to study the effects of light and defoliation on plant growth, plant carbon/nutrient balance, soil inorganic N content, and microbial activity and biomass in soil. Each microcosm contained three substrate layers – mineral soil, humus and plant litter – and one Nothofagus solandri var. cliffortioides seedling. The experiment constituted of the presence or absence of two treatments in a full factorial design: shading (50% decrease in light) and artificial defoliation (approximately 50% decrease in leaf area in the beginning of the growing season). At the end of the experiment a range of above‐ground and below‐ground properties were measured. The shading treatment reduced root and shoot mass, root/shoot ratio and leaf production of the seedlings, while the defoliation treatment significantly decreased leaf mass only. Leaf C and N content were not affected by either treatment. Shading increased NO ₃–N concentration and decreased microbial biomass in humus, while defoliation did not significantly affect inorganic N or microbes in humus. The results show that plant responses to above‐ground treatments have effects which propagate below ground, and that rather straightforward mechanisms may link above‐ground and below‐ground effects. The shading treatment, which reduced overall seedling growth and thus below‐ground N use and C allocation, also led to changes in humus N content and microbial biomass, whereas defoliation, which did not affect overall growth, did not influence these below‐ground properties. The study also shows the carbon/nutrient balance of N. solandri var. cliffortioides seedlings to be highly invariant to both shading and defoliation.     

Limited period of graviresponsiveness in germinating spores of Ceratopteris richardii

Rhizoids of the fern Ceratopteris richardii Brogn. usually emerge 40 h after germination is initiated by light, and more than 90% of them emerge growing in a downward direction. However, when the spores are germinated on a clinostat, the emerging rhizoids show no preferential orientation. This indicates that under normal 1 · g conditions the initial growth direction of rhizoids can be oriented by gravity. If the orientation of the spores is changed 3 h or less after the start of germination, the growth direction of most emerging rhizoids becomes downward relative to the new orientation. However, if the orientation of the spores is changed by 180° 8 h or more after germination is initiated by light, most rhizoids emerge growing upward; i.e., the same direction as if there had been no orientation change. Emerged rhizoids also do not change their direction of growth if their orientation is changed. These results indicate that the growth direction of emerging rhizoids is set by gravity prior to actual emergence, and that the time of full orientation responsiveness is limited to a period ranging from the initiation of germination to about 3–4 h after the start of germination. There is a gravity-oriented nuclear movement beginning at about 13 h after germination, and this movement appears to predict the initial growth direction of rhizoids.These studies were made possible by grant NAGW 1519 to S.J.R. and grant NGT-51065 to E.S.E., both from the National Aeronautics and Space Administration.     

Light partitioning among species and species replacement in early successional grasslands

Abstract. We studied canopy structure, shoot architecture and light harvesting efficiencies of the species (photon flux captured per unit above‐ground plant mass) in a series of exclosures of different age (up to 4.5 yr) in originally heavily grazed grassland in N Japan.Vegetation height and Leaf Area Index (LAI) increased in the series and Zoysia japonica, the dominant in the beginning, was replaced by the much taller Miscanthus sinensis. We showed how this displacement in dominance can be explained by inherent constraints on the above‐ground architecture of these two species. In all stands light capture of plants increased with their above‐ground biomass but taller species were not necessarily more efficient in light harvesting. Some subordinate species grew disproportionally large leaf areas and persisted in the shady undergrowth. Some other species first grew taller and managed to stay in the better‐lit parts of the canopy, but ultimately failed to match the height growth of their neighbours in this early successional series. Their light harvesting efficiencies declined and this probably led to their exclusion. By contrast, species that maintained their position high in the canopy managed to persist in the vegetation despite their relatively low light harvesting efficiencies. In the tallest stands ‘later successional’ species had higher light harvesting efficiencies for the same plant height than ‘early successional’ species which was mostly the result of the greater area to mass ratio (specific leaf area, SLA) of their leaves. This shows how plant stature, plasticity in above‐ground biomass partitioning, and architectural constraints determine the ability of plants to efficiently capture light, which helps to explain species replacement in this early successional series.     

Male orientation on vocalization perches could optimize acoustic signal transmission in anurans

Acoustic signals are distorted by vegetation, wind currents, or other sounds when transmitted through the environment. Consequently, vocalizations with features that optimize sound transmission or behaviors that improve the efficacy of communication have evolved in many animal species. Among behavioral strategies, some species call from perches above the ground to increase the propagation distance of their acoustic signals. However, the orientation in the perch also influences the transmission of the vocalizations, so that frogs calling from different orientations (i.e., horizontal, upward, or downward) may affect differently the quality and efficacy of sound transmission. We implemented a sound transmission experiment to test for the effect of calling orientation (upward, downward, and horizontal) and distance on the attenuation and degradation of advertisement calls in the common dink frog Diasporus diastema. We broadcasted and re‐recorded advertisement calls at 2 m height, setting the speaker in three directions (upward, downward, and horizontal) to simulate different signaler orientations. We found that attenuation of the advertisement calls is significantly reduced when the speaker was directed either upward or downward, rather than horizontally. However, the degradation of call is lower when the speaker is direct horizontally. Since calls produced from either upward or downward orientations could travel farther, they could be used to signal male spatial location, while calls produced from a horizontal position could provide information on male quality at shorter distances at advanced phases of courtship.     

A simulation model of production,seasonal changes in biomass and distribution of eelgrass (Zostera marina) in Lake Grevelingen

Summary A simulation model has been described, based on data from Lake Grevelingen, The Netherlands, as a predictive tool for lake management. The model has been developed as part of a large-scale aquatic modelling effort in Lake Grevelingen, carried out in close cooperation with the Delft Hydraulics Laboratory and the Delta Department, Environmental Research Division of the Ministry of Transport and Public Works. Available data on growth rates per unit eelgrass biomass, obtained with the leaf-marking technique, and on above- en below ground biomass and shoot density changes per unit area have been used. A space limitation depending on density of the above ground biomass and a growth limitation due to shortage in below ground biomass have been introduced. The seasonal changes in eelgrass production, both above and below ground, have been simulated as functions of the external forcing variables light, water temperature, wind generated water movements and of the internal control variables due to aging of the plant material. The vertical distribution of eelgrass can be partly explained from the modelling results on space, light and below ground biomass limitations. From the shore down to about 1 m waterdepth the above ground eelgrass biomass suffers from space limitation. Between 1 and 2 m production and biomass reach maximum values. Between 2 and 3 m waterdepth the above ground eelgrass growth is limited by the availability of below ground biomass. Between 3 and 5 m waterdepth both below ground biomass and light are the growth-limiting factors. Below 5 m waterdepth light is not sufficient to sustain net growth of eelgrass from rhizomes. Together with additional data — not used in the model — on seed production and growth of eelgrass shoots from seeds the vertical and horizontal distribution of the dominant macrophyte in the lake can be explained.     

Fruiting in Sphaerobolus with Special Reference to Light

Sphaerobolus grows and, provided there is sufficient illumination,fruits readily on oatmeal agar or on malt agar. No effect oflight on vegetative growth can be demonstrated. On the maltmedium, increased fruiting occurs with increase of nutrientup to 4 per cent, malt, but at higher concentrations fruitingis not increased and may be retarded. A chemically defined mediumwith starch as the carbon source allows fruiting, but at a lowlevel. Temperature has a profound effect on basidiocarp development;above 25 C. no fruit-bodies are normally formed although vegetativegrowth is approximately optimal at that temperature. For overallfruit-body production at 20 C, light above 100 lux is necessaryand light remains a limiting factor up to about 1, 000 lux.Under continuous light of suitable intensity, fruit-bodies continueto develop and discharge glebal-masses for many weeks. Thereis a distinct periodicity of discharge with (at 20 C.) about12 days between peaks of activity. This corresponds with thetime taken for basidiocarp initiation and development. A number of developmental stages of the basidiocarp are recognized.The final stage, glebal-mass discharge from stellately openedfruit-bodies, is indifferent to light, but all other stagesare stimulated by light. The light intensity for effective stimulationfalls during development and for the penultimate stage an intensityas low as 1 lux is effective. Only light of wave-length below500 mµ is active in overall basidiocarp development. Inthe sensitive region between 400 mµ and 500 mµ,there appear to be peaks of sensitivity around 440 mµand 480 mµ. In alternating light and darkness, simulating natural conditions,glebal-mass discharge occurs in the light periods. With a regimenof 24 hours light and 24 hours of darkness discharge is mainlyin the dark periods.     

The interstitial nucleus of Cajal of the cat. II. Effects of kainic acid lesion on vertical optokinetic nystagmus and after-nystagmus

The effects of bilateral lesions of the interstitial nucleus of Cajal (INC) by ka?nic acid on vertical optokinetic nystagmus (OKN) and after-nystagmus (OKAN) were studied in four cats: in three cats, in the acute stage from 1 to 60 days after the lesions; in the fourth cat, they were studied 3 years after the lesions were made. Histological control of lesions showed that the whole INC was bilaterally destroyed in two cats of the acute group and only the upper part of INC in the third cat. In the chronic cat, the density of cell bodies in both INC was lower than normal. In the acute group, the cats exhibited a spontaneous downward eye drift in light and in darkness. During an upward optokinetic stimulation, the effect of INC lesions was dramatic: upward slow phases and downward quick phases of OKN were abolished. Sixty days post lesions, small upward slow eye movements were again observed. During a downward optokinetic stimulation, the defect was much less; in particular, after a slight impairment of downward slow phases, during the first days post lesions, they recovered quickly. The secondary optokinetic after nystagmus (OKAN II) ensuing a downward OKN was cancelled and did not reappear 60 days post lesions. In the chronic stage, three years after the lesions, during an upward optokinetic stimulation, the cat showed upward slow phases with velocities close to normal. However, upward slow phases were curved: the velocity at the end of the slow phases was lower than at the beginning. After an upward OKN (the direction of slow phases gives the direction of the OKN and OKAN), the ensuing OKAN was present but abnormal.     

Adaptive longitudinal growth of first-order lateral roots of a woody species (Spartium junceum) to slope and different soil conditions—upward growth of surface roots

First-order lateral roots originating in the upper part of the taproot of a woody species, usually termed surface roots, grow close beneath the soil surface, even on irregular or sloping ground. In slope condition, in fact, the surface roots can assume upward as well as downward growth. Existing knowledge on the controls over root direction does not fully explain these field observations.

Two different soil types and sloping conditions were selected in field condition to explore the behaviour of the surface roots in the woody species Spartium junceum L. The root system 3D architecture was measured with a 3D digitizer and the angle of growth (0° = vertically downwards) and the radial direction (0° = horizontally downslope or northwards) of all root segments measured.

Surface roots were more numerous in clay soil than in loam soil, independently from the slope inclination. They had initial angles larger than 90°, i.e. they grew upwards only in clay soil. The subsequent angles of growth maintained this value only in steep-slope condition, showing a clear soil type x slope inclination interaction. The initial angle of all first-order lateral roots decreased linearly with depth of origin on the taproot always in relation to the soil type, with this relationship being stronger in clay soil.

These findings showed that the liminal angle (the preferred angle of growth) of surface roots was mainly affected by the soil type rather than the soil surface inclination. Thus, upward growth must stand in the plasticity of the plagiotropic response of these secondary laterals rather than in a strong internal control.     

Some effects of cultural treatments on virus diseases of cultivated mushroom, Agaricus bisporus

Inoculating crops of a white strain of mushroom with virus-infected cultures delayed cropping and decreased yields; the effects were progressively less the later that infection occurred, and/or the larger the amount of healthy spawn used. Crops in trays inoculated at a single site usually developed three zones: (1) a barren zone about the site of inoculation, progressively enlarging as the crops aged; and surrounded by (2) a band of stunted mushrooms bordering an outer area (3) of apparently healthy crop, in which a few sporophores might show hard-gill or watery-stipe abnormalities. Mycelial isolates taken at different distances from the sites of inoculation grew at different rates, growth being inversely proportional to the virus content of the fruit-bodies as estimated by electron microscopy and serology. Isolates from virus-free mushrooms grew rapidly on agar media, producing white fluffy colonies with many coarse strands, whereas those obtained from the few fruit-bodies near sites of inoculation were brown, adpressed to the medium, and grew very slowly. Between these extremes a continuous range of intermediates occurred, most of which remained constant when subcultured. Isolates taken at successive intervals from the same site in a tray had progressively smaller growth rates. Unsterilized mushroom-growing equipment (e.g. trays) is thought to carry viruses in infected spores and mycelial fragments which infect later crops without inducing characteristic zones. Instead, cropping of the whole tray declines with each successive flush. Such infection was prevented by heat-sterilizing the trays between crops.     

SEEDLING MORPHOLOGY IN MARAH (CUCURBITACEAE) RELATED TO THE CALIFORNIAN MEDITERRANEAN CLIMATE

Studies in reproductive ecology were made in indigenous, western American plants in the genus Marah (Cucurbitaceae), with particular attention given plants of M. oreganus occurring in the Berkeley Hills near San Francisco Bay in California. These tuberous perennials produce capsular fruits on their annual aboveground shoots; the fruits dehisce in early summer, each one exposing about three large seeds with an average seed weight of 1.05 g. The embryo of a M. oreganus seed has two thick and fleshy cotyledons packed with protein granules. The embryonic axis, with shoot and root apices, is ca. 0.5-1.0 mm long, roughly ½0 or less the length of the seed. In the Berkeley Hills dispersal of the seeds is accomplished by nocturnal rodents, after which germination begins with the fall rains and cooler temperatures of November and December. Instead of a radicle emerging first from the seed at germination, the minute radicle and epicotyl are pushed or carried far out of the seed, down into the soil, by the elongating bases of the cotyledons. These cotyledon bases, or petioles, are fused, and as they elongate they form a hollow tube that bears the embryonic axis at its extreme tip. The cotyledonary petiole tube ceases elongation by January, when it may be 5-25 or more cm long in a seedling of M. oreganus. Then, from its tip, the radicle grows downward and the epicotyl upward—up the hollow petiole tube. The green shoot (epicotyl) reaches the soil surface by early March in this area, completes the first season's growth, and dries up by late May, when the arid summer season is beginning. But even before the epicotyl grows out of the petiole tube and above ground, the seedling's hypocotyl begins to enlarge, forming a tuber. The fleshy cotyledon blades remain in the seed coat below ground, and some food from the blades is transferred to the tuber that produces shoots in the following growing seasons. This pattern of germination and seedling establishment is now known for species of Marah and for a very few other dicotyledonous plants, all of them growing mainly in areas of hot and dry habitat that are generally referred to as having Mediterranean climate. This elongation of the fused hypogeal cotyledons is considered a complex adaptation in dicotyledons that helps ensure fast and successful seedling establishment in seasonally arid areas such as “Mediterranean” California.     

Cultivation characteristics ofIsaria japonica

Cultivation characteristics of fruit-body (synnema) formation ofIsaria japonica were examined using liquid and solid media in order to produce fruit-bodies on a large scale. Mycelia grew well at 18–28°C on PDA medium with an initial pH of 7.0. The formation of fruit-bodies ofI. japonica was induced by lowering temperature to below 20°C in PD liquid medium. In sawdust-rice bran basal medium mixed with pupal powder prepared from silkworms (Bombyx mori), the fresh weight of fruit-bodies increased with increasing content of pupal powder. The highest yields of fruit-bodies were obtained in carbon-rich barley grain medium supplemented with pupal powder. The fruit-bodies grown under CO₂ concentrations of 1,000 μl/L had coral-like, many-branched synnemata with numerous conidiospores, whereas those formed under high concentrations (9,000 μl/L) of CO₂ had unbranched and longer synnemata. High concentrations of CO₂ remarkably inhibited conidiospore formation on synnemata. Continuous high-intensity illumination at 2.93 W·m⁻² inhibited the elongation of synnemata, and low-intensity illumination at 0.088 W·m⁻² slightly inhibited the branching of synnemata. Fruit-bodies were produced on the pupa metamorphosed from living larvae ofAgrotis fucosa placed on the surface of a culture ofI. japonica incubated in sawdust-rice bran medium.     

Phototropic Curvature in Phycomyces

The distribution of curvature and of bending speed along the cell's growing region are studied during steady state phototropic bending. At the start, elemental bending speed parallels the known axial distribution of growth rate. Hence regional phototropic sensitivity is initially determined by the local growth rate, and unilateral visible light acts proportionally at all levels of the growth zone. In the later course of bending, the bending speed distribution shifts downward instead of progressing upward in step with the cell's elongation. Furthermore, during phototropic inversion reversed bending begins high in the growth zone and progresses downward while normal bending continues below. These spatial and temporal changes in the distribution of differential growth are considered to be due to a fixed rate of supply of material used in growth that is transported from lower regions of the cell and asymmetrically distributed within the growth zone.     

Effects of light and low temperature on the reciprocal style curvature of Flexistylous Alpinia Species (Zingiberaceae)

The style curvature (flexistyly) of Alpinia species in ginger family is a unique plant organ movement because the style of each flower curves twice during its 1-day anthesis and styles of two phenotypes of each Alpinia species in the same population synchronously curve in opposite directions at the same time. In this study, we investigated the effects of low temperature and light conditions on these reciprocal style movements. Our results indicate that low temperature cannot change the direction of each curvature movement, but can slow down these movements and decrease the curve degrees. Light did not affect the upward curvature of the cataflexistylous morph, but the degrees of downward curvature decreased in darkness. For the anaflexistylous morph, the downward curvature only occurred in darkness, but curved directly upward in light condition; after the first (downward) curvature, the second (upward) movement only occurred in light, but did not occur if styles maintained in darkness. These results suggest that low temperature does not stimulate style curvature; light is the necessary condition for the upward movement of the anaflexistylous morph. The stimuli that induced curvature movements in the two morphs were different. Both two curvatures of the cataflexistylous style and downward movement of the anaflexistylous style were controlled via an endogenous program, while the upward movement of the anaflexistylous style was controlled by light.     

Evolution of complex life cycles in trophically transmitted helminths. I. Host incorporation and trophic ascent

Links between parasites and food webs are evolutionarily ancient but dynamic: life history theory provides insights into helminth complex life cycle origins. Most adult helminths benefit by sexual reproduction in vertebrates, often high up food chains, but direct infection is commonly constrained by a trophic vacuum between free‐living propagules and definitive hosts. Intermediate hosts fill this vacuum, facilitating transmission to definitive hosts. The central question concerns why sexual reproduction, and sometimes even larval growth, is suppressed in intermediate hosts, favouring growth arrest at larval maturity in intermediate hosts and reproductive suppression until transmission to definitive hosts? Increased longevity and higher growth in definitive hosts can generate selection for larger parasite body size and higher fecundity at sexual maturity. Life cycle length is increased by two evolutionary mechanisms, upward and downward incorporation, allowing simple (one‐host) cycles to become complex (multihost). In downward incorporation, an intermediate host is added below the definitive host: models suggest that downward incorporation probably evolves only after ecological or evolutionary perturbations create a trophic vacuum. In upward incorporation, a new definitive host is added above the original definitive host, which subsequently becomes an intermediate host, again maintained by the trophic vacuum: theory suggests that this is plausible even under constant ecological/evolutionary conditions. The final cycle is similar irrespective of its origin (upward or downward). Insights about host incorporation are best gained by linking comparative phylogenetic analyses (describing evolutionary history) with evolutionary models (examining selective forces). Ascent of host trophic levels and evolution of optimal host taxa ranges are discussed.     

辽东山区长白落叶松人工林天然更新障碍分析

通过对辽东山区不同间伐强度下长白落叶松林内种子库,种子萌发,幼苗出现、成活与生长的观测,分析了长白落叶松天然更新的主要影响因素.结果表明:40年生落叶松人工林的种子雨中,有30%的种子有活力,可以满足天然更新的需要;土壤种子库的种子主要分布在枯枝落叶层,4月下旬林内种子开始萌发,6月幼苗数量达到最大,幼苗累计出现率与间伐强度(光照)之间无显著相关,只受到地被物的影响;随间伐强度的增加,幼苗平均存活率提高,但林下幼苗生长缓慢,苗高均不超过6 cm;林内当年幼苗9月全部消失,没有超过1年生的幼苗,但皆伐迹地的更新幼苗存活较多、生长良好.初步确定地被物和光照是辽东山区长白落叶松天然更新的主要障碍因子.     

Development of appressoria on conidial germ tubes of <Emphasis Type="Italic">Erysiphe</Emphasis> species

Modes of branching of appressoria on conidial germ tubes of 36 Erysiphe spp. were studied. Only unlobed appressoria, termed alobatus pattern, were seen in E. lonicerae, E. magnifica and E. symphoricarpi. Viewed from above with light or scanning electron microscopes, other species had ± irregular lobing, but from below in the plane of contact with the substrate successive dichotomous branchings at 120° were seen to produce a five-lobed appressorium within 6 h. Each division produced a temporarily dormant outward-facing lobe and an inward limb that continued growth and division to form the axis of curved, hooked, single- or double-headed symmetrical or asymmetrical structures in a helicoid cyme-like pattern. Outlines of extracellular material after removal of germinated conidia confirmed this manner of branching. After 36 h some lobes re-divided forming botryose or jigsaw patterns even extending with extra appressoria to form candelabra-like structures. Conidia developed only one true germ tube; rarely secondary unswollen tubes emerged from spare shoulders or ends. The same true germ tubes developed initially on host surfaces, where secondary tubes and/or extensions from appressorial lobes grew into colony-forming hyphae. Lobed appressoria of Neoerysphe and Phyllactinia also branched at 120°. Podosphaera xanthii exhibited a simpler branching pattern.