Age-specific seasonal storage dynamics of<Emphasis Type="Italic">Phragmites australis</Emphasis> rhizomes: a preliminary study

Age-specific seasonal rhizome storage dynamics of a wetland stand of Phragmites australis (Cav.) Trin. ex Steud. in Japan, were investigated from April to October 2000. For each sampling date, above- and below-ground biomass and age-specific rhizome bulk density, ?_rhiz were measured. Seven rhizome age classes were recognized, from <1 year to six years old, based on their position within the branching hierarchy as main criteria and rhizome color, condition of nodal sheaths and condition of the shoots attached to vertical rhizomes as secondary criteria. P. australis stand was moderately productive, having a net aerial and below-ground production of 1980 and 1240 g m^?2, respectively, and a maximum mean shoot height of 2.33 ± 0.12 m. In spring, shoot growth started at the expense of rhizome reserves, decreasing the rhizome biomass as well as ?_rhiz. Both parameters reached the seasonal minimum in May followed by a subsequent increase, indicating a translocation of reserves to rhizomes from shoots after they become self supporting. For each sampling date, ?_rhiz increased with rhizome age. Given that the quantity of reserves remobilized by the rhizomes for spring shoot growth, as assessed by the drop in bulk density from April to May, were positively correlated (r = 0.97, P < 0.05) with rhizome age, it is proposed that for spring shoot formation older rhizomes remobilize stored reserves more actively than younger ones. Given that the accumulation of rhizome reserves (rise in bulk density) from May to August, May to September or May to November was negatively correlated (r = 0.97, 0.92 and 0.87, respectively, P < 0.05) with rhizome age, it seemed possible that younger rhizomes were ‘recharged’ at a higher rate than older ones. These resource allocation mechanisms pertaining seasonal rhizome storage dynamics are of paramount importance in formulating management and conservation strategies of wetlands and aquatic habitats. Our results indicate that a harvest of above-ground biomass from May to June would be more effective in reducing the growth than a harvest in July to August or later, when rhizome reserves have already been replenished. However, the latter may remove a larger shoot bound nutrient stock, still preserving a healthy stand for the subsequent years.     

Rhizome age structure of three populations ofPhragmites australis (Cav.) Trin. ex Steud.: Biomass and mineral nutrient concentrations

The rhizome age structure for populations ofPhragmites australis (Cav.) Trin. exSteud. was determined at three sites of the T?eboň Biosphere Reserve (Czech Republic). These sites were classified according to plant species composition and phosphorus availability as oligotrophic (Branná), mesotrophic (Ro?mberk East) and hypertrophic (Ro?mberk West).P.australis stands at these sites were expanding, stabilized and retreating, respectively. Rhizomes were sampled within the terrestrial parts of the reed stands (at a water depth of about 10 cm). The rhizomes were dated according to the position in the branching hierarchy and to supplementary morphological criteria. The analysis of the branching pattern revealed that at the sites studied, the rhizomes typically branch only once a year. The longest life span of rhizomes (six years) was found for the stable reed population at Ro?mberk East. The reduced rhizome life span at Ro?mberk West (four to five years), accompanied by a reduced branching frequency of horizontal rhizomes, was ascribed to a greater incidence of anaerobic conditions associated with the permanent flooding of the hypertrophic organic soil. The maximum life span of the Branná population (four years) probably corresponds to the total age of the reed population. The concentrations of nitrogen, phosphorus, and potassium in the tissue decreased with rhizome age while those of calcium and magnesium increased. The effect of rhizome age on mineral-nutrient concentrations was more pronounced than the effect of site on these concentrations. The mineral nutrient concentrations were not closely associated with reed performance.     

Colour choice behaviour in the pollen beetle Meligethes aeneus (Coleoptera: Nitidulidae)

The pollen beetle Meligethes aeneus Fabricius (Coleoptera, Nitidulidae), a pest of oilseed rape (Brassica napus), is known to respond to coloured stimuli; however, current understanding of the underlying mechanisms of colour choice in this species is limited. In the present study, physiological and behavioural experiments are conducted to determine the response of the pollen beetle to colours in the field. Spectral sensitivity is measured in 10 animals using the electroretinogram technique. Light flashes (100 ms) at varied wavelengths (340–650 nm, 10‐nm steps) and at different light intensities are applied to the eye after dark adaptation. In behavioural experiments in the field, 100 water traps of varying colours (from yellow to green to blue with varying amounts of white and black added, and with known spectral reflectance) are set out on a bare soil field in May 2008. The mean spectral sensitivity curve of M. aeneus peaks at 520 nm; however, a model template fitted to the long wavelength tail of the observed curve reveals a peak at approximately 540 nm (green). A secondary sensitivity peak is observed in the ultraviolet (UV) range (370 nm). A total of 2482 pollen beetles are captured in the coloured traps. The results show that the pollen beetles' preference for yellow over other colours can be modelled as a colour opponent mechanism (green versus blue); however, further experiments are needed to specify responses to colours with higher UV reflectance. These findings may be used to optimize trap colours for monitoring to help develop integrated pest management strategies for pollen beetle control.     

Geographic patterns in fruit colour diversity: do leaves constrain the colour of fleshy fruits?

We tested for geographic patterns in fruit colour diversity. Fruit colours are thought to promote detection by seed dispersers. Because seed dispersers differ in their spectral sensitivities, we predicted that fruit colour diversity would be higher in regions with higher seed disperser diversity (i.e. the tropics). We collected reflectance data on 232 fruiting plant species and their natural backgrounds in seven localities in Europe, North and South America, and analysed fruit colour diversity according to the visual system of birds—the primary consumer types of these fruits. We found no evidence that fruit colours are either more conspicuous or more diverse in tropical areas characterised by higher seed disperser diversity. Instead, fruit colour diversity was lowest in central Brazil, suggesting that fruit colours may be more diverse in temperate regions. Although we found little evidence for geographic variation in fruit hues, the spectral properties of fruits were positively associated with the spectral properties of backgrounds. This result implies that fruit colours may be influenced by selection on the reflectance properties of leaves, thus constraining the evolution of fruit colour. Overall, the results suggest that fruit colours in the tropics are neither more diverse nor more conspicuous than temperate fruits, and that fruit colours may be influenced by correlated selection on leaf reflectance properties. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The colour of flowers in spectrally variable illumination and insect pollinator vision

The spectral reflectance of differently coloured Australian native plant flowers and foliage was measured and plotted in a colour triangle to represent the colour space of the honeybee. Spectral variations in illumination are shown to significantly change plant colours for bee vision without colour constancy. A model of chromatic adaptation based upon the von Kries coefficient law shows a reduction in plant colour shift, with the degree of correction depending upon position in colour space. A set of artificial reflectances is used to map relative colour shift caused by spectrally variable illumination for the entire colour space of the honeybee. The rarity of some flower colours in nature shows a correlation to a larger colour shift for these colours when illuminated by spectrally variable radiation. The model of chromatic adaptation is applied to illuminations used in a behavioural study on honeybee colour constancy by Neumeyer 1981. Surface colours used by Neumeyer are plotted in colour space for the various illuminations. The results show that an illumination-dependent colour shift correlates to a decrease in the frequency of bees correctly choosing a colour to which it was trained. Accepted: 23 February 1998     

扎龙湿地不同生境芦苇种群根茎构件的年龄结构

芦苇是典型的长根茎型克隆植物,天然种群主要依靠根茎的营养繁殖进行更新。采用单位土体挖掘取样,按实际生活年限划分根茎龄级的方法,对扎龙湿地芦苇种群不同龄级根茎进行调查。结果表明,6月份,4个生境芦苇种群根茎构件均由2—6a的5个龄级组成,7—10月份均由1—6a的6个龄级组成。6—10月份,1a根茎长度比率逐渐增加;除盐碱生境略有增加外,其他3个生境2a、3a比率均小幅减少;4—6a比率逐渐减小,根茎长度比率均以3a最大,依次是2a,4a,1a,5a,最高的6a最小。1—3a根茎生物量比率逐渐增加;4—6a比率逐渐减小,均以3a根茎生物量比率最大,依次是4a,2a,5a,6a,以最低1a最小。不同龄级根茎长度和生物量比率与返青后实际生长时间之间均较好地符合直线函数关系(R~20.91,P0.01;R~20.81,P0.05),6—9月份,根茎长度呈衰退型年龄结构,10月份又转为稳定型结构,整个生长季根茎生物量均为衰退型结构。不同龄级根茎构件在芦苇种群中的地位和作用不同,根茎构件的年龄结构蕴含着种群调节的重要信息。     

Predicting the Effectiveness of <Emphasis Type="Italic">Phragmites</Emphasis> Control Measures using a Rhizome Growth Potential Bioassay

In the last century, Phragmites australis (common reed) has expanded from a minor component of the mid-Atlantic tidal wetlands to a dominant species in many locations. Expansion of Phragmites results in decreased plant diversity and alterations to the tidal characteristics of the marsh, resulting in decreased wetland value. Management efforts have used a variety of strategies in an attempt to control its expansion. We tested a greenhouse bioassay that provided insight into the rhizome vitality of six herbicide-treated sites in the Alloway Creek Watershed, NJ well in advance of the growing season. At three sites, rhizomes were exhumed and classified by depth (0–25 cm and 25–75 cm) and appearance (color and firmness). Concurrently, the same protocol was followed, but conducted on an areal basis at three additional sites. Material was grown in sand under greenhouse conditions void of nutrient supplements for 70 days, after which shoots were removed and the rhizomes replanted for 30 days. Effectiveness of control strategies was quantified by examining rhizome color, vitality, and shoot densities in the field. Color was indicative of quality of rhizome reserves. Less than 0.2% of the firm, brown rhizomes produced shoots upon initial planting and none produced shoots upon replanting, whereas 50.9% of white rhizomes produced shoots on initial planting. Rhizome vitality was quantified by examining shoot emergence and the morphology of the shoots. Coupling rhizome vitality with observed field densities resulted in a predictive capability, and shoot density and biomass predictions were compared to field measurements in July 2001. We tested and accurately predicted the relative shoot densities and shoot biomass of the three sites for which we collected rhizome material on an areal basis. The result is a rapid, valuable, and cost-effective monitoring tool that can quickly quantify the effects of past control methods and predict future growth potential.     

扎龙湿地芦苇种群不同龄级根茎构件的季节动态

克隆植物根茎具有营养繁殖和扩展种群的功能,也是芽和分株生理整合的通道。根茎构件具有出生、死亡及年龄等种群统计特征,不同龄级根茎的季节动态可以反映根茎的存活和衰老过程。采用单位土体挖掘取样,对扎龙湿地4个生境芦苇种群根茎构件进行野外调查,比较不同龄级根茎长度、生物量和干物质贮量的季节动态。结果表明:7—10月份,1a根茎长度、生物量和干物质贮量均呈指数函数增加,在生长季中后期有一个持续时间较长的生长和物质积累时期。6—10月份,2a、3a根茎长度呈线性函数增加,4—6a根茎长度呈线性函数减少;2—4a根茎生物量和2—5a根茎干物质贮量呈二次函数先减少后增加,5a、6a根茎生物量和6a根茎干物质贮量呈幂函数减少。整个生长期内,根茎长度和根茎生物量均以3a最大,根茎长度以最高的6a最小,根茎生物量以最低的1a最小;根茎干物质储量以5a最大,以最低的1a最小。4个生境芦苇种群根茎长度、生物量和干物质贮量在龄级间的差异及差异序位稳定,在新根茎的产生、老根茎的存活以及根茎寿命与养分消耗和储藏上均具有稳定的生物学特性,不同龄级根茎在种群中的地位和作用以及对种群的贡献不同。     

Broad spectral sensitivities in the honeybee's photoreceptors limit colour constancy

Colour constancy allows for visual systems to be view stimuli independent of changes in spectral illumination. Chromatic adaptation is likely to be an important mechanism in colour constancy and can be explained by use of the von Kries coefficient law. Chromatic adaptation is compared for the honeybee and three hypothetical visual systems. It is shown that the spectral breadth and asymmetry of photoreceptors in the honeybee may limit colour constancy. In particular, it is demonstrated that the absorption of short-wavelength radiation by the cis-band of chromophore is responsible for a poorer correction for bee colours rich in ultraviolet reflectance. The results are discussed in relation to theoretical considerations of von Kries colour constancy and the physiology of eye design in some other species for which colour constancy has been demonstrated. Accepted: 14 August 1999     

Molecular cytogenetics of Androctonus scorpions: an oasis of calm in the turbulent karyotype evolution of the diverse family Buthidae

After years of qualitative and subjective study, quantitative colour science is now enabling rapid measurement, analysis and comparison of colour traits. However, it has not been determined how many replicates one needs to accurately quantify a species' colours for studies aimed at broad cross‐species trait comparison. We address this major methodological knowledge gap. We first quantified and assessed the variance in colour within and between species. Reflectance spectra of flowers from ten plant species and plumage of 20 bird species were measured using a spectrometer, and reflectance (i.e. brightness) and tetrahedral colour‐space coordinates were calculated. analysis of variance (ANOVA) analyses indicate that there is far more variation in the colours of birds and flowers between species (> 77%) than within species. A Mean Absolute Deviation from the Mean test was applied to indicate the sampling replication required for each species. Tetrahedral coordinates were sampled precisely with only one individual per species. Greater replication was needed to sample reflectance with the desired precision, particularly for darker coloured species. Our findings will allow researchers to allocate their sampling effort in a way that maximises the precision of their colour data collection. The fact that only a few replicates per species are necessary will greatly facilitate broad cross‐species comparisons of colour in the future. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 114 , 69–81.     

扎龙湿地不同生境芦苇种群根茎数量特征及动态

采用单位土体取样,计测长度和生物量的调查与统计方法,对扎龙湿地保护区4个生境芦苇种群根茎数量特征进行比较分析。结果表明,芦苇5月10日左右返青后进入营养生长期,根茎长度6—8月份缓慢增加,8—10月份显著增加,后期是前期的3.5—10.3倍,生长季中后期是种群新根茎补充和生长的主要时期,不仅实现了种群空间扩展,并为营养繁殖储备更多繁殖芽;根茎生物量和干物质贮量6—8月份逐渐减少,8—10月份又逐渐增加,均以生长季末期的10月份最大,并均显著地(P0.05)高于其他月份,种群根茎养分的消耗主要供给根茎芽的萌发和幼株生长,根茎养分的储藏又为翌年种群的更新及扩展提供物质保障,种群对地下根茎存在明显的养分"超补偿性"贮藏现象。种群根茎长度和生物量均以湿生生境最大,依次为旱生生境、水生生境,盐碱生境最小,根茎干物质贮量以旱生生境最大,依次为湿生生境、水生生境,盐碱生境最小。种群根茎长度与返青后实际生长时间之间均较好地符合直线函数关系,种群根茎生物量和干物质贮量与生长时间之间较好地符合二次曲线函数关系,R2在0.804—0.997之间,拟合方程均达到了显著或极显著(P0.01)水平。4个生境芦苇种群在根茎长度、生物量、干物质贮量等数量特征均表现出由遗传因素控制的比较稳定的季节动态规律,在生境间的差异及其差异序位又均基本稳定,均表现出明显的土壤因子环境效应,其中土壤含水量、有机质、速效氮为正向驱动,p H、速效磷为负向驱动,土壤含水量、p H对根茎数量特征的驱动作用更突出。     

Not so colourful after all: eggshell pigments constrain avian eggshell colour space

Birds'' eggshells are renowned for their striking colours and varied patterns. Although often considered exceptionally diverse, we report that avian eggshell coloration, sampled here across the full phylogenetic diversity of birds, occupies only 0.08–0.10% of the avian perceivable colour space. The concentrations of the two known tetrapyrrole eggshell pigments (protoporphyrin and biliverdin) are generally poor predictors of colour, both intra- and interspecifically. Here, we show that the constrained diversity of eggshell coloration can be accurately predicted by colour mixing models based on the relative contribution of both pigments and we demonstrate that the models'' predictions can be improved by accounting for the reflectance of the eggshell''s calcium carbonate matrix. The establishment of these proximate links between pigmentation and colour will enable future tests of hypotheses on the functions of perceived avian eggshell colours that depend on eggshell chemistry. More generally, colour mixing models are not limited to avian eggshell colours but apply to any natural colour. Our approach illustrates how modelling can aid the understanding of constraints on phenotypic diversity.     

Phragmites australis at an extreme altitude: Rhizome architecture and its modelling

In central EuropePhragmites australis is a lowland plant, occurring rarely up to the tree line. In the Velká Kotlina cirque (Jeseníky mountains, NE Czech Republic), where it reaches its maximum altitude at about 1350 m a.s.l., its culms are 0.5–0.7 m high and the plants flower only in some years. During the last decade no germinable seeds have been observed. The architecture ofPhragmites rhizomes from this site was studied on seven randomly selected clonal fragments. They consisted of 3 to 10 partial tussocks (clumps) and 4 to 17 green shoots. The total length of the rhizomes was 9.7 to 50 m per plant. The number of nodes per plant was 96 to 431 and the longest internodes were 83 mm long. The number of side branches was 31 to 105 per plant. The branching angle depended on the type of branched rhizome. The mean angles of horizontal rhizomes, which connect individual tussocks, were relatively wide (modus 45°, arithmetic mean 37°), whereas within a tussock much sharper angles of branching prevailed (modal value 5°, arithmetic mean 15°). The mean internode-to-internode angle on continuing rhizomes was about 8°, with a wide variation. An architectural, spatially-explicit model ofPhragmites rhizome growth has been developed, showing that thePhragmites population in the studied locality can be maintained by vegetative multiplication, and seedling recruitment is not needed for its long-term persistence.     

Effects of rhizome age on the decomposition rate of Phragmites australis rhizomes

The change in dried rhizome samples that were left to decompose was investigated to elucidate the effects of rhizome age on the decomposition rate of Phragmites australis. Rhizomes were classified into five age categories and placed 30 cm below the soil surface of a reed stand. After 369 days of decay, new (i.e., aged less than one year) rhizomes had lost 84% of their original dry mass, compared with a loss of 41–62% for that of older rhizomes. The exponential decay rates of older rhizomes were nearly identical to that of aboveground biomass. The nitrogen (N) concentration increased to two times its original values, but the phosphorus (P) concentration remained constant after an initial loss by leaching. The carbon to nitrogen (C:N) and carbon to phosphorus ratios (C:P) leveled out at 22:1 to 38:1 and 828:1 to 1431:1, respectively, regardless of rhizome age. The results are important to understand the nutrient cycles of reed-dominant marsh ecosystems.     

Seasonality of carotenoid‐based plumage coloration: modelling wavelength‐specific change through spectral reconstruction

Plumage coloration has provided important model systems for research on signal expression. Whilst it had previously been assumed that moulting provided the only mechanism to change plumage coloration, recent studies have shown plumage colours to be seasonally dynamic, with implications both for the quantification of expression and for any signalling role. However, the mechanistic processes underlying such change remain uncertain. Here, we describe within‐moult shifts in expression of a carotenoid‐based colour trait – the yellow ventral plumage of the great tit Parus major – over a nine‐month timespan. We report that plumage chromaticity (‘colour’) – but not achromaticity (‘brightness’) – exhibits a marked seasonal decline, independent of sex, age or body condition, and at a constant rate across twelve environmentally heterogeneous plots within our study site. To gain a greater understanding of the mechanisms underlying this change we employed a spectral reconstruction approach, that generates predicted spectra for any timepoint within the sampling period. By comparing spectra for both early and late in the moult we show that the seasonal decline in chromaticity is driven by both a marked reduction in ultraviolet (UV) reflectance and, to a lesser extent, loss of active carotenoid pigments. Thus, our study shows that seasonal loss of chromaticity in the great tit is driven by altered reflectance primarily in the UV section of the spectrum, a finding made possible by the use of spectral compartmentalisation and multi‐parallel modelling to produce reconstructed spectra. Whether change in plumage coloration influences signal function will depend on the dynamics of the signalling system but it could clearly inflate patterns such as assortative mating and should be considered in studies of colour expression.     

Effect of water depth on the rhizome dynamics of Typha angustifolia

The rhizome dynamics of Typha angustifolia in relation to water depth were investigated. Observations were made in two different stands, namely Akigase from April 2002 to December 2003, and Teganuma from April 2003 to December 2004. The mean rhizome length in the two stands was not significantly different but the mean rhizome diameter and biomass showed a significant difference. The larger rhizome diameter in Teganuma is seemingly an important adaptation to maintain effective aeration of the rhizome by reducing the internal resistance; while larger biomass helps survival under anoxia for longer periods of time, it also provides strong anchorage to the substrate.     

Regeneration capacity and carbohydrate reserves in a clonal plant Rumex alpinus: effect of burial

The morphological and anatomical responses to different depths of burial were examined in Rumex alpinus (Polygonaceae), a perennial plant with monopodial, horizontally growing rhizome. Its segments, which consist of 12–20 internodes, 1 to 2 mm in length each, are products of single growing seasons. The rhizomes regenerated from 5, 10 and 20 cm, but failed to emerge from 30 cm. Number of internodes produced during a growing season was not affected by burial but the length of internodes increased up to about 30-fold. The rhizomes growing up to the surface were subsidized by older rhizome segments. In the case of deeply buried rhizomes the carbohydrate reserves of the last-year-segment were nearly completely depleted. Evolutionary significance of the regeneration capacity is discussed.     

Decomposition of aboveground and belowground organs of wild rice (Zizania latifolia): mass loss and nutrient changes

Decomposition of aboveground and belowground organs of the emergent macrophyte Z.latifolia was investigated using a litterbag technique for a period of 359 days in a freshwater marsh in Japan. Aboveground parts were classified into: leaves, sheaths and stems. Belowground parts were classified into: horizontal rhizomes (new rhizome, hard rhizome, soft rhizome) and vertical rhizome (stembase). The decay rate (k) was 0.0036 day⁻¹, 0.0033 day⁻¹ and 0.0021 day⁻¹ for leaves, sheaths and stems, respectively. For belowground parts, the decay rate varied considerably from 0.0018 day⁻¹ to 0.0079 day⁻¹, according to differences in the initial chemical compositions of rhizomes. After 359 days of decay, new rhizomes lost 94% of their original dry mass, compared with a loss of 48–84% for the other rhizomes. There was a significant positive relationship between litter quality and decay rate for horizontal rhizomes. For the new rhizomes, which had an internal nitrogen content of 24.2 mg N g⁻¹ dry mass, the mass loss was 40% higher than that of soft rhizomes, which had an internal N content of 9.8 mg N g⁻¹ dry mass. Over the period of 359 days, the nitrogen concentration in all rhizome types decreased to levels lower than initial values, but the phosphorus concentration remained constant after an initial leaching loss. Most nitrogen and phosphorus were lost during the first 45 days of decay. Changes in carbon to nitrogen (C:N) and carbon to phosphorus (C:P) ratios basically followed inversed trends of the nitrogen and phosphorus concentrations.     

Feather colours of live birds and museum specimens look similar when viewed by seabirds

Bird plumage and skin colour can be assessed from museum specimens. To determine whether these accurately represent the colours of live birds when viewed by birds themselves, we analysed the spectral reflectances of live and up to 100‐year‐old museum specimens of five seabird species (White‐faced Petrel Pelagodroma marina, Common Diving Petrel Pelecanoides urinatrix, Grey‐faced Petrel Pterodroma gouldi, Little Shearwater Puffinus assimilis and Fluttering Shearwater Puffinus gavia). Live birds had brighter colours than museum specimens, but there were no significant differences in the wavelengths reflected. Modelling indicated that seabirds would be able to detect colour changes in the skin, but not the feathers, of museum specimens, but only for species with blue or pink feet (Pelecanoides urinatrix and Puffinus assimilis). For seabirds, museum specimens are adequate proxies for feather colour but not for skin colour.