Electronic simulation of ganglion cells of generalized vertebrate cone retina

Electronic simulation of generalized vertebrate cone retina consists of 43x41 grid of red-, green-, and blue-sensitive cones. Each retinal element is simulated by a linear summator in series with a leaky integrator and spatial-temporal properties are developed by spatial organization of cone mosaic into unit hexagons and interplay of antagonistic inputs of differing time courses. Model has full compliments of horizontal and bipolar cells including color- and noncolor coding as well as single- and double-opponent receptive fields for bipolar cells. Electronic simulation also has negative feedback from L-horizontal cells to cones. Ganglion cells are formed by convergence of 7 bipolar cells, either all same and thus homogeneous, or else with a central-DPBC (or HPBC) and 6 surround-HPBCs (or DPBCs) and thus non-homogeneous. Responses of color- and non-color-coded ganglion cells as well as single- and double-opponents are investigated with stationary and moving light spots using white and colored lights. While responses to stationary light spots are predictable from digital models, responses to moving spots are complicated by differing time lags of components involved in total response. Therefore, responses to moving stimuli are more readily simulated by analogue models.     

Dynamics of L-type bipolar and phasic amacrine cells in the vertebrate cone retina

The model of the cone-L-HC circuit of the catfish retina (Siminoff 1985a) is extended to Luminosity bipolar cells (BC) and non-linear phasic amacrine cells (AC), but now applicable to the generalized vertebrate cone retina that involves only one cone type. Two types of BC's are simulated by linear transformation of 2 antagonistic inputs of differing time courses; the faster center field hyperpolarization from the cone and the slower surround field depolarization from the L-HC. The phasic AC was made non-linear by various methods: full- or half-wave rectification using either both or only one of the BC's as the inputs with rectification first and then summation or summation first and then rectification. A method is described using Laplace transforms in conjunction with the convolution theorem to obtain the impulse responses of BC's and AC's, in spite of the non-linearities of the AC even when used as feedback to the BC's. Since the input to the BC consists of 2 antagonistic inputs, feedback from the AC reeinforces one input and attenuates the other.     

Intermodulation components of the visual evoked potential: Responses to lateral and superimposed stimuli

Nonlinear interactions in the human visual system were studied using visual evoked potentials (VEPs). In one experiment (superimposed condition), all segments of a dartboard pattern were contrast reversed in time by a sum of two sinusoidal signals. In a second experiment (lateral condition), segments in some regions of the dartboard pattern were contrast reversed by a single sinusoid of one frequency, while segments in other (contiguous) regions of the pattern were contrast reversed by a single sinusoid of another frequency. An identical set of ten frequency pairs was used in each experiment. The frequency pairs were chosen such that the difference between frequencies in each pair was 2 Hz. Amplitudes and phases of the sum and difference frequency components of the VEP (intermodulation terms) were retrieved by Fourier analysis and served as measures of nonlinear interactions. The use of input pairs with a fixed separation in frequency enabled the estimation of the temporal characteristics of the visual pathways prior to a second linear stage. The use of superimposed and lateral conditions revealed antagonistic contributions to the VEP, possibly reflecting direct-through excitatory and lateral inhibitory pathways, respectively.Supported by grants from the U.S. National Eye Institute, the Esther A. and Joseph Klingenstein Fund, and the Harry Frank Guggenheim Foundation     

Chromatic induction: Responses of neurophysiological double opponent units?

Equations have been derived that improve the quantification of sensory equidistant colour and lightness differences. This has been achieved by a physiological approach involving non-linear responses of cone mechanisms and two subsequent stages of linear opponent transformation to describe the Munsell System (Seim and Valberg, 1980). Using the formulation for the first opponent stage, colours induced into an achromatic center field by a chromatic surround varying in purity, are shown to follow the same power function of the opponent coordinates for all hues. By analogy, a physiological model for colour coding and colour induction is offered. Double opponent neurones with spatially antagonistic, spectrally opponent and symmetric receptive fields constitute the units of the model. Colour induction is related to lateral excitation and colour differences to response differences of these units.     

Aridity drives spatiotemporal patterns of masting across the latitudinal range of a dryland conifer

Masting, or the synchronous and irregular production of seed crops, is controlled by environmental conditions and resource budgets. Increasing temperatures and shifting precipitation regimes may alter the frequency and magnitude of masting, especially in species that experience chronic resource stress. Yet the effects of a changing climate on seed production are unlikely to be uniform across populations, particularly those that span broad abiotic gradients. In this study, we assessed the spatiotemporal patterns of masting across the latitudinal distribution of a widely distributed dryland conifer species, piñon pine Pinus edulis. We quantified seed cone production from 2004 to 2017 using cone abscission scars in 187 trees from 28 sites along an 1100 km latitudinal gradient to investigate the spatiotemporal drivers of seed cone production and synchrony across populations. Populations from chronically hot and dry areas (greater climatic water deficits and less monsoonal precipitation) tended to have greater interannual variability in seed cone production and smaller crop sizes. Mast years generally followed years with low vapor pressure deficits and high precipitation during key periods of the reproductive process, but the strength of these relationships varied across the region. Populations that received greater monsoonal precipitation were less sensitive to late summer vapor pressure deficits during seed cone initiation yet more sensitive to spring vapor pressure deficits during pollination. Spatially correlated patterns of vapor pressure deficit better predicted synchrony in seed cone production than geographic distance, and these patterns were conserved at distances up to 500 km. These results demonstrate that aridity drives spatiotemporal variability in seed cone production. As a result, projected increases in aridity are likely to decrease the frequency and magnitude of masting in these dry forests and woodlands. Declines in seed production may compound climatic limitations to recruitment and impede tree regeneration, with cascading effects for numerous wildlife species.     

Colour vision in the passeriform bird,Leiothrix lutea: correlation of visual pigment absorbance and oil droplet transmission with spectral sensitivity

The visual receptors in the retina of the passeriform bird Leiothrix lutea were examined microspectro-photometrically. The rods had a maximum absorbance close to 500 nm. Four spectrally different classes of single cone were identified with typical combinations of photopigments and oil droplets: a long-wave sensitive cone with a photopigment P568 and a droplet with a cut-off wavelength at 564 nm, a middle-wave sensitive cone with a P499 and a droplet with a cut-off at 506 nm, a short-wave sensitive cone with a P454 and a droplet with maximum absorbance below 410nm and an ultraviolet sensitive cone with a P355 and a transparent droplet. Double cones possessed a P568 in both the principal and accessory members. A pale droplet with variable absorbance (maximal at about 420 nm) was associated with the principal member whereas the ellipsoid region of the accessory member contained only low concentrations of carotenoid. The effective spectral sensitivities of the different cone classes were calculated from the characteristic combinations of oil droplets and photopigments and corrected for the absorbance of the ocular media. Comparison of these results with the behavioural spectral sensitivity function of Leiothrix lutea suggests that the increment threshold photopic spectral sensitivity of this avian species is mediated by the 4 single cone classes modified by neural opponent mechanisms.Abbreviations LWS long wave sensitive - MWS middle wave sensitive - SWS short wave sensitive (cones)     

Development of Partial Ontogenic Resistance to Powdery Mildew in Hop Cones and Its Management Implications

Knowledge of processes leading to crop damage is central to devising rational approaches to disease management. Multiple experiments established that infection of hop cones by Podosphaera macularis was most severe if inoculation occurred within 15 to 21 days after bloom. This period of infection was associated with the most pronounced reductions in alpha acids, cone color, and accelerated maturation of cones. Susceptibility of cones to powdery mildew decreased progressively after the transition from bloom to cone development, although complete immunity to the disease failed to develop. Maturation of cone tissues was associated with multiple significant affects on the pathogen manifested as reduced germination of conidia, diminished frequency of penetration of bracts, lengthening of the latent period, and decreased sporulation. Cones challenged with P. macularis in juvenile developmental stages also led to greater frequency of colonization by a complex of saprophytic, secondary fungi. Since no developmental stage of cones was immune to powdery mildew, the incidence of powdery mildew continued to increase over time and exceeded 86% by late summer. In field experiments with a moderately susceptible cultivar, the incidence of cones with powdery mildew was statistically similar when fungicide applications were made season-long or targeted only to the juvenile stages of cone development. These studies establish that partial ontogenic resistance develops in hop cones and may influence multiple phases of the infection process and pathogen reproduction. The results further reinforce the concept that the efficacy of a fungicide program may depend largely on timing of a small number of sprays during a relatively brief period of cone development. However in practice, targeting fungicide and other management tactics to periods of enhanced juvenile susceptibility may be complicated by a high degree of asynchrony in cone development and other factors that are situation-dependent.     

Modelling Mediterranean pasture dynamies

Pasture vegetation in an open woodland of Quercus rotundifolia subjected to periodic ploughing was sampled in spring during 8 consecutive years. The frequency of herbaceous species was recorded in a total of 69 permanent plots located on 5 adjacent sites with similar lithology, slope and orientation but differing in age since previous ploughing.Vegetation dynamics expressed as trajectories of permanent plots in a non-metric multidimensional scaling space has been modelled in terms of evironmental variables. By fitting a generalized linear model, the dynamics are shown to be related to years since last ploughing, geographical location of plots, total annual rainfall and November rainfall. Meteorological patterns of the sampling period are also described.Abbreviations GLM = Generalized Linear Model - NMDS = Non-metric Multi-Dimensional Scaling - UPGMA = Unweighted Pair-Group Method using Arithmetic Averages     

Variations in the short wavelength cut-off of the solar UV spectra.

Cloud and solar zenith angle (SZA) are two major factors that influence the magnitude of the biologically damaging UV (UVBD) irradiances for humans. However, the effect on the short wavelength cut-off due to SZA and due to clouds has not been investigated for biologically damaging UV for cataracts. This research aims to investigate the influence of cloud and SZA on the short wavelength cut-off of the spectral UVBD for cataracts. The spectral biologically damaging UV for cataracts on a horizontal plane was calculated by weighting the spectral UV measured with a spectroradiometer with the action spectrum for the induction of cataracts in a porcine lens. The UV spectra were obtained on an unshaded plane at a latitude of 29.5 degrees S. The cut-off wavelength (lambdac) was defined as the wavelength at which the biologically damaging spectral irradiance was 0.1% of the maximum biologically damaging irradiance for that scan. For the all sky conditions, the short wavelength cut-off ranged by 12 nm for the SZA range of 5 to 80 degrees and the maximum in the spectral UVBD ranged by 15 nm. Similarly, for the cloud free cases, the short wavelength cut-off ranged by 9 nm for the same SZA range. Although, cloud has a large influence on the magnitude of the biologically damaging UV for cataracts, the influence of cloud on the short wavelength cut-off for the biologically damaging UV for cataracts is less than the influence of the solar zenith angle.     

马铃薯立枯丝核菌拮抗菌QHZ11的实时荧光定量PCR快速检测与应用

【背景】植物根际促生菌(plant growth-promoting rhizobacteria,PGPR)在根际的定殖是其发挥作用的基础,直观有效的跟踪技术和定量方法是研究PGPR在根际原位分布规律的重要工具。【目的】建立一种马铃薯黑痣病病原菌——立枯丝核菌拮抗菌QHZ11的实时荧光定量PCR快速检测体系,并检测拮抗菌QHZ11在马铃薯根际的动态变化。【方法】根据GenBank中登录的类芽孢杆菌及近源菌株gyrB基因序列差异筛选特异性引物,优化反应条件;通过盆栽试验对马铃薯根际拮抗菌进行快速检测。盆栽试验设3个处理,T1:对照(无菌水,CK);T2:QHZ11菌悬液灌土(QHZ11);T3:将功能菌在有机肥中进行二次固体发酵制成生物有机肥(BOF11)。【结果】筛选出拮抗菌QHZ11的专用引物为gyrB-F/gyrB-R;建立的拮抗菌QHZ11实时荧光定量PCR检测方法特异性好、灵敏度高且重复性较好,线性相关系数为0.999 8,检测组内变异系数均在1%以内,扩增效率为0.9,可检测出1×103-1×1010copies/g-soil的拮抗菌,具有检出限低和扩增效率高的特点。盆栽试验...     

Electronic simulation of cones,horizontal cells and bipolar cells of generalized vertebrate cone retina

Electronic analogue of my theoretical model of generalized vertebrate cone retina [Siminoff: J. Theor. Biol. 86, 763 (1980)] is presented. Cone mosaic is simulated by 25x21 grid of phototransistors that have colored filters mounted in front of then to produce red-, green-, and blue-sensitive cones arranged in a trichromatic retina. Each retinal element is simulated by Summator-Integrator and unit gain voltage invertes are used to give correct polarities to output voltages. Dynamic properties of retinal elements are developed solely by temporal interplay of antagonistic input voltages with differing time courses, and spatial organization of receptive fields is developed by unit hexagons that precisely define cone input voltages to subsequent elements. Electronic model contains both color- and non-colorcoded channels. Negative feedback from L-horizontal cells to cones, electrical coupling of like-cones, and electrical coupling of like-horizontal cells are simulated by feedfoward circuits. Stray light is present due to light scattering properties of colored filters used to simulate color selectivety of cones. Stationary and moving spots of white and colored lights of varied sizes and intensities are used to study characteristics of electronic analogue. Results demonstrate practicality of electronic simulation to function analogous to real cone retinas to process visual stimuli and give information to higher centers as to size, shape, color and motion of objects in visual world.     

Evolution of pleiotropic alleles for maturation and size as a consequence of predation

Understanding life-history evolution requires knowledge about genetic interactions, physiological mechanisms and the nature of selection. For platyfish, Xiphophorus maculatus, extensive information is available about genetic and physiological mechanisms influencing life-history traits. In particular, alleles at the pituitary locus have large and antagonistic effects on age and size at sexual maturation. To examine how predation affects the evolution of these antagonistic traits, I examined pituitary allele evolution in experimental populations differing in predation risk. A smaller size, earlier maturation allele increased in frequency when predators were absent, while a larger size, later maturation allele increased in frequency when predators were present. Thus, predation favours alleles for larger size, at the cost of later maturation and reproduction. These findings are interesting for several reasons. First, predation is often predicted to favour early reproduction at smaller sizes. Second, few studies have shown how selection acts on alleles that affect age and size at sexual maturation. Finally, many studies assume that trade-offs between these life-history traits result from antagonistic pleiotropy, with alleles that positively affect one trait negatively affecting another, yet this is rarely known. This study unequivocally demonstrates that genetically based trade-offs affect life-history evolution in platyfish.     

Effect of secondary flow on biological experiments in the cone-plate viscometer: methods for estimating collision frequency, wall shear stress and inter-particle interactions in non-linear flow.

We present a theoretical analysis of fluid flow and particle interactions in the cone-plate viscometer under conditions typically applied in biological studies. The analysis demonstrates that at higher shear rates, besides linear primary flow in the rotational direction, prominent non-linear secondary flow causes additional fluid circulation in the radial direction. Two parameters, the cone angle and Reynolds number, characterize flow in the viscometer over all ranges of shear rate. Our results indicate that secondary flow causes positional variations in: (i) the velocity gradient, (ii) the direction and magnitude of the wall shear stress at the plate surface, (iii) inter-particle collision frequency, (iv) magnitude and periodicity of normal and shear forces applied during particle-particle interactions, and (v) inter-particle attachment times. Thus, secondary flow may significantly influence cellular aggregation, platelet activation and endothelial cell mechanotransduction measurements. Besides cone-plate viscometers, this analysis methodology can also be extended to other experimental systems with complex non-linear flows.     

Spouse-to-Spouse Transmission and Evolution of Hypervariable Region 1 and 5’ Untranslated Region of Hepatitis C Virus Analyzed by Next-Generation Sequencing

Hepatitis C virus (HCV) transmission between spouses remains poorly characterized, largely due to the limited availability of samples from the early stage of infection, as well as methodological constraints. A fifty-eight year-old male developed acute hepatitis C infection and his 53-year old spouse has been HCV-positive for over 10 years. Serum samples were collected from both at the time of acute hepatitis C diagnosis in male (baseline) and then at 9 and 13 months. Hypervariable region 1 (HVR1) and 5’ untranslated region (5’UTR) sequences were amplified and subjected to next generation sequencing (NGS) using a pyrosequencing platform. Genetic variants were inferred by Shorah reconstruction method and compared by phylogenetic and sequence diversity analysis. As the sequencing error of the procedure was previously determined to be ≤ 1.5%, the analysis was conducted with and without the 1.5% cut-off with regard to the frequency of variants. No identical HVR1 variants were identified in spouses at baseline and follow-up samples regardless whether the cut-off was applied or not. However, there was high similarity (98.3%) between a minor baseline donor variant (1.7% frequency) and the most abundant baseline recipient variant (62.5% frequency). Furthermore, donor and recipient strains clustered together when compared to 10 control subjects from the same area and infected with the same HCV subtype. There was an increase in HVR1 complexity (number of genetic variants) over time in both spouses. In contrast, the 5''UTR region was stable and of low complexity throughout the study. In conclusion, intrafamilial HCV transmission may be established by a very minor variant and investigation of this phenomenon requires high-sensitivity assays, such as NGS.     

The apparent mass of the human body exposed to non-orthogonal horizontal vibration

Apparent masses of 15 male and 15 female subjects have been measured during exposure to various directions of horizontal vibration. Twenty vibration conditions were used in the experiment. In each of five directions (0, 22.5, 45, 67.5 and 90° to the mid-sagittal plane) subjects were exposed to random vibration in the frequency range of 1.5–20 Hz at 0.25, 0.5 and 1.0 m s⁻² r.m.s. The five remaining conditions were selected to give measurements whereby the magnitude of the x-component of the vibration was fixed and the y-component changed and vice-versa. Two peaks were observed in the apparent masses. The first peak occurred at about 3 Hz and reduced in frequency with increases in vibration magnitude. The frequency of the first peak also reduced as the direction of vibration changed from 0 to 90°. The magnitude of the peak increased as the vibration magnitude and direction increased. The second peak occurred at about 5 Hz and decreased in both frequency and magnitude with increases in vibration magnitude. There was no change in the frequency of the second peak with vibration direction, although the magnitude of the peak decreased as the angle of vibration to the mid-sagittal plane increased. Increasing the magnitude of the x-component of vibration whilst using a fixed y-component changed the magnitude of the first peak but did not change the frequency of the first or any characteristics of the second peak. In contrast, increasing the y-component of vibration whilst using a fixed x-component changed the frequencies and magnitudes of both peaks. Predictions of the response at 45° by applying the principle of superposition to data measured at 0 and 90° showed that the response of the body with direction was not linear. This implies that the apparent mass in non-orthogonal axes cannot be predicted from the apparent masses measured in orthogonal directions.     

Two models of the recognition and detection of texture-defined letters compared

We quantified texture segregation by measuring psychophysically the percentage correct detection scores for each of a set of 10 texture-defined (TD) letters using the temporal two-alternative forced choice method, and at the same time quantified spatial discrimination of the TD form of measuring psychophysically the percentage correct letter recognition scores for the 10 letters. Ten levels of task difficulty were created by adding noise dots to the texture patterns. The resulting psychophysical data were used to test and compare models of the detection and recognition of texture-defined letters. Each model comprised a sequence of physiologically plausible stages in early visual processing. Each had the same first, second and third stages, namely linear orientation-tuned spatial filters followed by rectification and smoothing. Model 1 had only one non-linear stage. Model 2 had two non-linear stages. In model 2 the second non-linear stage was cross-orientation inhibition. This second non-linear stage enhanced the texture borders by, in effect, comparing textures at different locations in the texture pattern. In both models, the last stage modelled either letter detection or letter recognition. Letter recognition was modelled as follows. We passed a given letter stimulus through the first several stages of a model and, in 10 separate calculations, cross-correlated the output with a template of each of the 10 letters. From these 10 correlations we obtained a predicted percentage correct letter recognition score for the given letter stimulus. The predicted recognition scores closely agreed with the experimental data at all 10 levels of task difficulty for model 2, but not for model 1. We conclude that a borderenhancing algorithm is necessary to model letter recognition. The letter-detection algorithm modelled detection of part of a letter (a single letter stroke) in terms of the signal-to-noise ratio of a letter-segment detector. The predicted letter detection scores fitted the data closely for both models.     

Effects of low-pass filter combinations on lower extremity joint moments in distance running

Inverse dynamics is a standard tool in biomechanics, which requires low-pass filtering of external force and kinematic signals. Unmatched filtering procedures are reported to affect joint moment amplitudes in high impact movements, like landing or cutting, but are also common in the analysis of distance running. We analyzed the effects of cut-off frequencies in 94 rearfoot runners at a speed of 3.5 m/s. Additionally, we investigated whether the evaluation of footwear interventions is affected by the choice of cut-off frequencies. We performed 3D inverse dynamics for the hip, knee and ankle joints using different low-pass filter cut-off frequency combinations for a recursive fourth-order Butterworth filter. We observed fluctuations of joint moment curves in the first half of stance, which were most pronounced for the most unmatched cut-off frequency combination (kinematics: 10 Hz; ground reaction forces (GRFs): 100 Hz) and for more proximal joints. Peak sagittal plane hip joint moments were altered by 94% on average. We observed a change in the ranking of subjects based on joint moment amplitude. We found significant (p < 0.001) footwear by cut-off frequency combination interaction effects for most peak joint moments. These findings highlight the importance of cut-off frequency choice in the analysis of joint moments and the assessment of footwear interventions in distance running. Based on our results, we propose to use matched cut-off frequencies around 20 Hz in order to avoid large artificial fluctuations in joint moment curves while at the same time avoiding a severe removal of physiological high-frequency signal content from the GRF signals.     

Adaptive radiation through phenological shift: the importance of the temporal niche in species diversification

Abstract 1. Phenological shift in oviposition in seed predators may be a key factor for adaptive radiation if temporal differences lead to less intense competition. 2. This hypothesis was tested at two sites in the French Alps in three sympatric species of larch cone flies grouped into two phenological groups (early and late) differing in adult emergence and oviposition timing by approximately 2 weeks. The present study assessed the intensity of competition within and between groups by measuring four larval traits. Cone traits were measured, and the impact of early species parasitism on cone development was assessed. 3. The occupation of the central axis of a developing cone by one early larva has a strong detrimental effect on cone growth and seed production. However, there was almost no correlation between the variables measured on the cones and on the larvae, suggesting that the resources available were not limiting. 4. Inter‐group competition had no significant effect on early larvae. In contrast, both inter‐ and intra‐group competition had a significant negative effect on late larvae length (–11% and –16% respectively), dry mass (–8% and –23%), and lipid mass (–15% and –26%). The intensity of competition was stronger among larvae in the same phenological group, which is consistent with the hypothesis that shifts in oviposition promote adaptive radiation in larch cone flies by reducing competition among larvae.     

A linear chromatic mechanism drives the pupillary response.

Previous studies have shown that a chromatic mechanism can drive pupil responses. The aim of this research was to clarify whether a linear or nonlinear chromatic mechanism drives pupillary responses by using test stimuli of various colours that are defined in cone contrast space. The pupil and accommodation responses evoked by these test stimuli were continuously and simultaneously objectively measured by photorefraction. The results with isochromatic and isoluminant stimuli showed that the accommodative level remained approximately constant (< 0.25 D change in mean level) even when the concurrent pupillary response was large (ca. 0.30 mm). The pupillary response to an isoluminant grating was sustained, delayed (by ca. 60 ms) and larger in amplitude than that for a isochromatic uniform stimulus, which supports previous work suggesting that the chromatic mechanism contributes to the pupillary response. In a second experiment, selected chromatic test gratings were used and isoresponse contours in cone contrast space were obtained. The results showed that the isoresponse contour in cone contrast space is well described (r(2) = 0.99) by a straight line with a positive slope. The results indicate that a /L - M/ linear chromatic mechanism, whereby a signal from the long wavelength cone is subtracted from that of the middle wavelength cone and vice versa, drives pupillary responses.     

Gender performance in a cultivated cohort of the cycad Zamia integrifolia (Zamiaceae)

A progeny of the native Florida cycad Zamia integrifolia grown from seeds planted in 1986 was monitored until 1995 to record mortality and the nature and time of expression of primary and secondary sex characters. In addition to gender-specific cone morphologies, males and females differed in secondary sex characters such as age at first cone production, frequency of cone production, mean cone numbers in second and later coning episodes, and, in older plants, mean leaf and branch numbers. Gender differences expressed themselves at different stages in the life history: their nature and extent varied during the years following sexual maturation. By 1995, 46% of the plants in the progeny had died, most of them before producing cones. Prior to 1988 the mean leaf number of plants that died did not differ from that of survivors, but the mean leaf number of plants dying between 1988 and 1989 was 0.4 times that of the survivors during that period, suggesting reduced vigor prior to death. Mean age at first cone production was 5.8 yr for males and 6.6 yr for females. Mean dry masses of individual male cones increased between the first and second coning episodes, but not between the second and third coning episodes. Mean dry masses of the entire cone crop of individual males increased through the third coning episode due to an increase in mean cone number per episode, but mean cone number was unchanged between the third and fourth coning episodes. Mean dry mass of unpollinated female cones did not change between the first and second coning episodes; mean cone numbers did not change between the first and third coning episodes. After the first coning episode, males produced higher mean cone numbers than females. By 1995, the mean dry mass of an individual male's cone crop was greater than that of a female. Coning frequency of males was 1.7 times greater than that of unpollinated females, suggesting a gender difference in the genetic control of coning frequency. Coning frequency of females pollinated 1 or 2 yr previously was reduced compared with that of unpollinated females. Cone production did not affect subsequent leaf production by either gender. Mean leaf numbers increased in some years and not in others. Mean leaf numbers of males and females did not differ prior to cone production. After cone production mean leaf numbers of males were greater than of females. Mean age of males producing first branches was 6.3 yr, with a mean of 2.5 first branches per plant. Mean age of females producing first branches was 7.7 yr, with a mean of 2.5 first branches per plant. By 1995 the mean branch number of males was 5.7 per plant and of females was 2.7 per plant. Between 1993 and 1995 the mean branch number of males and females increased incrementally, but mean leaf numbers did not change. In early years of branching, leaf number increased with branch number; higher mean leaf numbers of males of an age class thus reflected their earlier branching. Males produced first cones earlier than females. Since branch production was associated with cone production, higher branch numbers of males in an age class reflected their earlier first cone production. In 1995 the sex ratio of known males and females in the progeny was 1:1, with a few individuals not having produced cones by that year.