Seasonal changes in photosynthesis of four understory herbs in deciduous forests

Seasonal patterns of photosynthesis and respiration of single leaves of four understory perennial herbs in deciduous forests were investigated in relation to their leaf growth and light conditions on the forest floor.Anemone flaccida shows rapid growth of leaf area and high rates of gross photosynthesis at light saturation (Psat) in its early stage of development. Its photosynthetic activity is restricted to a brief period of high light intensity before the closure of overstory canopies.Disporum smilacinum possesses light-photosynthesis curves of the shade-leaf type throughout its whole growing period. A shading experiment has shown that this plant is low-light adapted and can utilize weak light efficiently. The light-photosynthesis curve ofSyneilesis palmata shifts from the sun-leaf type to the shade-leaf type in response to the seasonal change of light regime on the forest floor. Evergreen leaves ofPyrola japonica have three year longevity, and light-photosynthesis curves of the shade-leaf type. They maintain some photosynthetic activity even in late autumn and winter.     

PHOTOSYNTHESIS AND CARBON ALLOCATION IN TIPULARIA DISCOLOR (ORCHIDACEAE), A WINTERGREEN UNDERSTORY HERB

Seasonal patterns of photosynthesis and carbon allocation were determined for Tipularia discolor, a summer-deciduous wintergreen orchid of the southeastern United States, to assess the effects of environmental conditions and leaf age on carbon acquisition and allocation patterns. There was no shift in the optimum temperature for photosynthesis (T_opt) on a seasonal basis and T_opt (≈26 C) was at least 10 C higher than daily maximum air temperature during most of the growing season. Lack of photosynthetic adjustment in Tipularia to seasonal fluctuations in temperature and light suggested that the photosynthetic characteristics of this wintergreen were more similar to those of spring ephemerals than to those of evergreens and summer-active herbs. The decline in photosynthetic capacity during the winter growing season for Tipularia, largely due to leaf age effects, gradually reduced net photosynthetic rates in the field despite more favorable light and temperature conditions. Photosynthesis in the field was primarily limited by environmental conditions in early- and mid-season and by photosynthetic capacity in late-season. A ¹⁴CO₂ labelling experiment demonstrated that patterns of carbon allocation to vegetative structures were affected by the season of photosynthetic carbon fixation, whereas reproductive structures received 21% of the recovered labelled carbon regardless of the period of labelling. Carbon acquired and stored during all periods of the growing season was used to produce new vegetative and reproductive structures.     

PIGMENT AND PHOTOSYNTHETIC RESPONSES OF OSCILLATORIA AGARDHII (CYANOPHYTA) TO PHOTON FLUX DENSITY AND SPECTRAL QUALITY1

The effects of photon flux density (PFD) and spectral quality on biomass, pigment content and composition, and the photosynthetic activity of Oscillatoria agardhii Gomont were investigated in steady-state populations. For alterations of PFD, chemostat populations were exposed to 50, 130 and 230 μmol photons·m^?2·s^?1 of photosynthetic active radiation (PAR). Decreases in biomass, chlorophyll a (Chl a) and c-phycocyanin (CPC) contents, and CPC: Chl a and CPC: carotenoid content was not altered. Increases in the relative abundances of myxoxanthophyll and zeaxanthin and deceases in the relative abundances of echinenone and β-carotene within the carotenoid pigments coincided with increasing PFD. Increases in Chl a-specific photosynthetic rates and maxima and decreases in biomass-specific photosynthetic rates and maxima with increasing PFD were attributed to increased light harvesting by carotenoids per unit Chl a and reduction in total pigment content, respectively. Responses to spectral quality were tested by exposing chemostat populations to a gradient of spectral transmissions at 50 μmol photons·m^?2·s^?1 PAR. Biomass differences among populations were likely attributable to the distinct absorption of the PAR spectrum by Chl a, CPC, and carotenoids. Although pigment contents were not altered by spectral quality, relative abundances of zeaxanthin and echinenone in the carotenoid pigments increased in populations exposed to high-wavelength PAR. The population adapted to green light possessed a greater photosynthetic maximum than populations adapted to other spectral qualities.     

不同种源油松幼苗的光合色素和非结构性碳水化合物对模拟氮沉降的短期响应

以不同种源(内蒙NM、北京BJ和山西SX)的3年生油松幼苗为研究对象,研究不同种源油松幼苗的光合色素以及非结构性碳水化合物(NSC)对氮沉降增加的短期响应。实验中设置5个氮处理:CK(0 kg hm-2a-1)、N1(15 kg hm-2a-1)、N2(25 kg hm-2a-1)、N3(50 kg hm-2a-1)、N4(150 kg hm-2a-1)。研究结果表明:(1)不同生长季,3个种源油松幼苗的光合色素对氮沉降增加的响应存在差异,但是只有BJ种源油松幼苗的叶绿素含量在生长季中期受到了氮沉降增加的显著促进作用,并在N4水平下达到最大值。(2)氮沉降的增加促进了3个种源油松幼苗NSC的转移和消耗,在生长季初期和中期,随着氮沉降水平的升高,3个种源油松幼苗的可溶性糖(SS)含量、淀粉(ST)含量以及总非结构性碳水化物(TNC)含量呈不同程度的降低。生长季末期,3个种源油松幼苗的SS和TNC明显积累。N4水平抑制了NM种源油松幼苗SS和BJ种源油松幼苗ST的累积,促进了NM种源油松幼苗ST含量和BJ种源油松幼苗SS含量的提高。氮沉降的增加显著抑制了SX种源油松幼苗NSC的积累,延长了幼苗的生长期,推迟了幼苗进入休眠的时间。     

Responses of leafing phenology and photosynthesis to soil warming in forest-floor plants

Phenological and physiological responses of plants to climate change are key issues to understand the global change impact on ecosystems. To evaluate the species-specific responses, a soil-warming experiment was conducted for seven understory species having various leaf habits in a deciduous forest, northern Japan; one evergreen shrub, one semi-evergreen fern, one summer-deciduous shrub, and four summer-green herbs. Soil temperature in the warming plots was electrically maintained 5 °C higher than control plots. Responses of leafing phenology highly varied among species: new leaf emergence of the evergreen shrub was delayed; senescence of overwintering leaves of the semi-evergreen fern was accelerated resulting in the shift to deciduousness; leaf shedding of the summer-deciduous shrub was accelerated. Among four summer-green species, only an earliest leaf-out species advanced growth initiation, but the period of growth season was not changed. Physiological responses to soil warming were also highly species-specific: the warming treatment increased the photosynthetic activity of the summer-deciduous shrub and one summer-green species, decreased that of the semi-evergreen fern, while other species did not show any changes in photosynthetic traits. Totally, the soil warming impacts on understory plants was apparent in spring. It was suggested that modification of snow conditions is important issue especially for plants with overwintering leaves. Responses of understory vegetation to climate change may highly vary depending on the composition of leaf habits in the cool-temperate forests.     

Seasonal changes in the photosynthetic response ofMercurialis perennis plants from different light regime conditions

Curves relating net photosynthetic rate to irradiance [P(I) curve relation] were estimated and analysed inMercurialis perennis L. plants stemming from three forest (spruce, beech and ash) stands with different tree leaf canopy development and different light regime. The saturating irradiance (I_s) reached the highest values in plants of all three stands in spring (spruce forest: 438 W m⁻², beech forest: 440 W m⁻² and ash forest: 367 W m⁻²), it declined sharply in the middle of the growing season (283, 285 and 297 W m^-2, respectively) and this I_s level persisted until autumn. A pronounced dynamics in plants from spruce and beech forests made itself manifest also in the adaptation (I_a) and compensating (I_c) irradiances, respectively. After a sudden decline in summer, values in autumn were close to those of the vernal season. The most pronounced parameter, which optimally expressed the adaptation ofMercurialis perennis to various light conditions, was the photosynthetic efficiency (α) calculated as the slope of the linear part of the curve relating net photosynthetic rate to irradiance. At the time of the highest P_{N sat.} value in course of the growing season (August) (spruce forest: 100, beech forest: 98.7 and ash forest: 85.8 μg CO₂ m⁻² s⁻¹), R_D was in its minimum; in autumn P_{N sat.} reached the lowest values which corresponded to the most intensive R_D. It was found thatMercurialis perennis plants stemming from forest stands with different light conditions did not make use equally of the altering light conditions in the course of the growing season. By the underlying analysis of P(I) curves this rhizomatous perennial herb (geophyte) may be characterized as a shade tolerant species.     

The effect of iron chelate fertilization of poplar upon CO2-uptake,leaf size,and content of leaf pigments and iron

Summary Potted poplars (strainsmarilandica, serotina andFlachslanden ofPopulus euramericana) which developed iron-deficiency symptoms (chlorosis of upper leaves, winter die-back of leader, flushing of lateral buds) were treated with a soil application of iron chelate to study the effect of iron nutrition upon CO₂-uptake, iron and pigment content of leaves, and leaf size of a tree species. Foliar content of each iron, chlorophyll, -carotene, lutein, and violaxanthin was significantly increased by the treatment. Chlorophyll b proved to be particularly sensitive to iron supply and the Q^a/b was also significantly altered.CO₂-uptake increased in fertilized and non-fertilized leaves with increasing light up to 40,000 Lux, but fertilized leaves assimilated more CO₂ than non-fertilized leaves, especially at light intensities from 5,000 Lux upwards. The assimilatory number was decreased by the iron application since larger amounts of chlorophyll were present in fertilized leaves. If CO₂-uptake was based upon an area unit basis the fertilizer effect became distinct even at 500 Lux. Thus CO₂-uptake is a quick, valuable measure of fertilizer responses.In severe cases, iron deficiency also affects leaf size and thus indirectly reduces photosynthetic activity. A chelate application during the growing season will not affect the size of leaves already formed but may considerably increase the size of leaves formed subsequent to the treatment.     

Is the Lower Shade Tolerance of Scots Pine,Relative to Pedunculate Oak,Related to the Composition of Photosynthetic Pigments?

Foliage of Scots pine (Pinus sylvestris L.) and pedunculate oak (Quercus robur L.) was collected in a mixed pine/oak forest at canopy positions differing in radiation environment. In both species, chlorophyll (Chl) a/b ratios were higher in foliage of canopy positions exposed to higher irradiance as compared to more shaded crown layers. Throughout the growing season, pine needles exhibited significantly lower Chl a/b ratios than oak leaves acclimated to a similar photon availability. Hence, pine needles showed shade-type pigment characteristics relative to foliage of oak. At a given radiation environment, pine needles tended to contain more neoxanthin and lutein per unit of Chl than oak leaves. The differences in pigment composition between foliage of pine and oak can be explained by a higher ratio of outer antennae Chl to core complex Chl in needles of P. sylvestris which enhances the efficiency of photon capture under limiting irradiance. The shade-type pigment composition of pine relative to oak foliage could have been due to a reduced mesophyll internal photon exposure of chloroplasts in needles of Scots pine, resulting from their xeromorphic anatomy. Hence, the higher drought tolerance of pine needles could be achieved at the expense of shade tolerance.     

Stemflow nutrient inputs to soil in a successional hardwood forest

Stemflow and throughfall from a regenerating (8-year-old) southern Appalachian hardwood forest were collected to examine the relative importance of tree bole nutrient leaching in response to acid deposition. Samples from nine (2 m²) stemflow collection plots were analyzed for four dormant season and 11 growing season rainstorm events. Results showed that, relative to throughfall fluxes, stemflow accounted, on average, for approximately 8.5% of total water reaching the forest floor during both dormant and growing season storms. Relative to foliar leaching, K-, SO₄-, and PO₄ ions appear to be the most easily leached ions from young tree stems. Proportional nitrate and base cation stemflow fluxes increased significantly (p<0.05) with growing-season storm-event duration, suggesting that the stemsurface nutrient pool is depleted by precipitation more slowly than the foliar pool. On average, proportional stemflow fluxes of SO₄ (12%) and K (14%) were consistently higher than reported maximum values for more mature forest stands, which indicates that small-scale stemflow inputs of ions such as these to the forest floor may be important in early successional ecosystems.     

A comparison of the responses of two tropical specialist herbivores to host plant patch size

Summary The effects of host plant patch size on the abundances of two specialist herbivores (the chrysomelid beetle, Acalymma innubum and the pentatomid bug, Piezosternum subulatum) were investigated in a natural forest community in the Virgin Islands. Abundances were compared early and late in the season in different sized patches of the cucurbit host plant (Cayaponia americana) growing in open habitat (with no surrounding plant community) and forest habitat (with diverse surrounding plant community). For both herbivore species, adult abundances per patch were positively correlated with patch leaf area, but there was a significant patch size effect (i.e., correlation between herbivore density per unit plant and patch leaf area) only for beetles in the forest habitat. Both herbivore species were significantly affected by surrounding plant diversity, but in opposite ways: beetles were more abundant in open patches whereas bugs were more abundant in forest patches. Relationships between abundance and patch size in open and forest patches changed through the season for both herbivore species. These changing abundance patterns are discussed with respect to (1) increases in the diversity of the plant community surrounding host plant patches, and (2) differences in herbivore movement patterns.     

Forest floor photosynthesis and respiration in a drained peatland forest in southern Finland

Background: Although forest floor forms a large biomass pool in forested peatlands, little is known about its role in ecosystem carbon (C) dynamics.

Aim: We aimed to quantify forest floor photosynthesis (P _FF) and respiration (R _FF) as a part of overall C dynamics in a drained peatland forest in southern Finland.

Methods: We measured net forest floor CO₂ exchange with closed chambers and reconstructed seasonal CO₂ exchange in the prevailing plant communities.

Results: The vegetation was a mosaic of plant communities that differed in CO₂ exchange dynamics. The reconstructed growing season P _FF was highest in the Sphagnum community and lowest in the feather moss communities. On the contrary, R _FF was highest in the feather moss communities and lowest in the Sphagnum community. CO₂ assimilated by the forest floor was 20–30% of the total CO₂ assimilated by the forest. The forest floor was a net CO₂ source to the atmosphere, because respiration from ground vegetation, tree roots and decomposition of soil organic matter exceeded the photosynthesis of ground vegetation.

Conclusions: Tree stand dominates C fluxes in drained peatland forests. However, forest floor vegetation can have a noticeable role in the C cycle of peatlands drained for forestry. Similarly to natural mires, Sphagnum moss-dominated communities were the most efficient assimilators of C.     

A hemiparasite in the forest understorey: photosynthetic performance and carbon balance of Melampyrum pratense

• Melampyrum pratense is an annual root‐hemiparasitic plant growing mostly in forest understorey, an environment with unstable light conditions. While photosynthetic responses of autotrophic plants to variable light conditions are in general well understood, light responses of root hemiparasites have not been investigated.
• We carried out gas exchange measurements (light response and photosynthetic induction curves) to assess the photosynthetic performance of M. pratense in spring and summer. These data and recorded light dynamics data were subsequently used to model carbon balance of the hemiparasite throughout the entire growth season.
• Summer leaves had significantly lower rates of saturated photosynthesis and dark respiration than spring leaves, a pattern expected to reflect the difference between sun‐ and shade‐adapted leaves. However, even the summer leaves of the hemiparasite exhibited a higher rate of light‐saturated photosynthesis than reported in non‐parasitic understorey herbs. This is likely related to its annual life history, rare among other understorey herbs. The carbon balance model considering photosynthetic induction still indicated insufficient autotrophic carbon gain for seed production in the summer months due to limited light availability and substantial carbon loss through dark respiration.
• The results point to potentially high importance of heterotrophic carbon acquisition in M. pratense, which could be of at least comparable importance as in other mixotrophic plants growing in forests – mistletoes and partial mycoheterotrophs. It is remarkable that despite apparent evolutionary pressure towards improved carbon acquisition from the host, M. pratense retains efficient photosynthesis and high transpiration rate, the ecophysiological traits typical of related root hemiparasites in the Orobanchaceae.

     

Seasonal patterns of acid fluctuations and resource storage in the arborescent cactus Opuntia excelsa in relation to light availability and size

Summary We investigated relationships between light availability, diel acid fluctuation, and resource storage in the arborescent cactus Opuntia excelsa growing in western Mexico. We compared canopy and understory individuals from a deciduous forest as well as open-grown plants of the same approximate size as those in the understory. During the wet season light availability and daily fluctuations in titratable acidity (an index of carbon uptake) were lower in the understory than in unshaded habitats. In the dry season all plants had reduced levels of acid fluctuation, with the smallest individuals, regardless of habitat, showing the greatest reduction. These data suggest that light availability in the forest understory constrains carbon assimilation during the wet season, but that a factor associated with plant size, possibly water status, limits carbon gain during the dry season. Plants in all habitats remained physiologically active for at least five months into the dry season. We suggest that this was possible due to the maintenance of constant concentrations of water and nitrogen in the photosynthetically active chlorenchyma. Parenchyma in terminal cladodes showed a different seasonal pattern of resource storage; water content and nitrogen concentration were reduced from the wet to the dry season in the parenchyma. Using the parenchyma to supply photosynthetic tissues during times of reduced resource availability allows O. excelsa to assimilate carbon during times of the year when most other trees in the forest are leafless.     

CHARACTERIZATION OF OCEANIC PHOTOSYNTHETIC PICOEUKARYOTES BY FLOW CYTOMETRY1

To interpret flow cytometric data that are routinely obtained on natural oceanic communities, 23 strains of photosynthetic picoeukaryotes belonging to four classes (Prasinophyceae, Chlorophyceae, Pelagophyceae, and Prymnesiophyceae) and six pigment types were investigated for their light scattering in the forward and right-angle directions, chlorophyll fluorescence, and DNA content as measured by flow cytometry. Cell she was assessed by Coulter counter, and pigment composition was measured by reverse-phase high-performance liquid chromatography. The size and GC% of the nuclear genome of cultured picoeukaryotes was measured from the fluorescence of DNA-specific dyes. Using these two parameters, we could discriminate species within pigment groups. DNA staining of preserved natural samples may also prove useful in discriminating cooccurring populations in situ as long as the communities are not too complex. Using the relationships that we established between size and light-scattering properties of the cells, we estimated equivalent diameters of picoeukaryotes in natural populations to be between 1.3 and 2 μm. Chlorophyll a content was between 6 and 16 fg·cel^?1 as calculated from relationships that we established between chlorophyll a content and red fluorescence of the cultured strains. With respect to size, chlorophyll a content, and pigment composition, Pelagomonas sp. strains (Pelagophyceae) appeared to be the most representative of the natural communities in subtropical ocean waters. In contrast, green coccoid strains, which often outcompete other strains in culture, might only be minor contributors to these communities.     

Adaptation of epiphytic bryophytes in the understorey attributing to the correlations and trade-offs between functional traits

This study explores adaptive strategies of epiphytic bryophytes in the understorey by investigating the photosynthetic characteristics, pigment concentrations and nutrient stoichiometry, as well as other functional traits of three trunk-dwelling bryophytes in a subtropical montane cloud forest in SW China. The results showed that their light-saturated net photosynthetic rate (A_nmax?L), light saturation point (I_sat), light compensation point (I_c) and dark respiration rate (R_d) were ca 0.55, 106.72, 4.17 and 0.25?μmol?m^?2?s^?1, respectively. Furthermore, the samples demonstrated photosynthetic down-regulation under high irradiance. These photosynthetic characteristics can be explained by higher total chlorophyll concentrations, specific leaf area, chlorophyll per unit leaf N (Chl/N), lower ratio of chlorophyll a to chlorophyll b (Chl a/b) and photosynthetic nitrogen-use efficiency. We suggest that the bryophytes adapted to the shaded understorey microhabitats through a series of correlations and trade-offs between functional traits.     

不同生境下濒危植物膝柄木幼树的生态适应性

膝柄木是我国极度濒危植物,也是广西滨海过渡带天然植被的重要组成树种.为了解光因子对膝柄木天然更新的限制影响,该文对林缘、林窗、林下三种不同光照生境下膝柄木幼树的生理和生长指标的年际变化特征进行了研究.结果表明:(1)光合有效辐射不足影响了膝柄木幼树的生长.林下幼树的地径、株高和叶面积增长量显著降低,而生长于光照充足林缘...     

Structure and dynamics of Japanese beech (Fagus japonica Maxim.) stools and sprouts in the regeneration of the natural forests

Structure and spatial distribution of stools and root-collar sprouts of Japanese beech (Fagus japonica) were studied to clarify the regeneration processes of the stool and the population, and the ecological importance of this stool formation in five quadrats of the natural forests with different forest floor vegetation on the Pacific side of Japan. F. japonica dominates in the canopy of each quadrat. Most of sprouts formed a circle around the root-collar and lowest parts of the parent stems of the stool with the youngest sprouts at the periphery. The regeneration by seedlings was slight especially on the forest floor vegetation of the dwarf bamboo Sasa. The variety of size structure of stems and the existence of dead traces and/or dead center in each stool, the apparent difference in stool size, and the aggregations of stools in the forests suggest that stool expansion and long persistence of the stool at a given location may contribute to compensate for the scarcity of regeneration by seedlings inhibited by dwarf bamboo, and by the other shrubs and herbs.     

SEASONAL PHOTOSYNTHETIC PERFORMANCE OF MACROCYSTIS INTEGRIFOLIA (PHAEOPHYCEAE)1

The seasonal photosynthetic performances of three age classes of blades of Macrocystis integrifolia Bory were estimated by studying their photosynthetic rate vs. irradiance curves and pigment contents for 15 months. All blade types were irradiance-saturated between 25 and 70 μE · m^?2· S^?1. Young and mature blade tissues had higher photosynthetic maxima and initial slopes on an area basis than older blade tissue. The latter, however, had pigment concentrations similar to those in mature blade tissues. All these parameters varied on a seasonal basis. The photosynthetic maxima ranged from 0.1–0.8 μmol · C · cm^?2· h^?t and showed two peaks, one in late summer-early fall and the other in late winter. Changes in initial slope and pigment concentrations in the blade tissues suggest that, changes in the size or efficiency of electron transfer in the photosynthetic unit occur. These data are discussed in relation to changes in seawater temperature and nitrate concentrations.     

Effects of flower and seed predators and pollinators on fruit production in two sequentially flowering congeners

We examined spatial and temporal variability of understory herbaceousvegetation on opposing north- and south-facing slopes in an eastern deciduousold-growth forest in southeastern Ohio, USA. Secondly, we explored theinfluenceof sampling scale and analytical technique on our assessment of diversitypatterns. The influence of aspect and seasonality were examined at varyingsampling scales using observed richness, evenness, andH diversity measures, non-parametric richnessestimators, species-area curves, and SHE analysis. Herb layer composition,abundance, and diversity were strongly influenced by location (north slope vs.south slope), seasonal sampling period (April, June, August), and plot size(micro (2 m²)- vs. meso (70m²)-scale samples). Although north and south plots werecompositionally distinct, they followed similar courses of change through thegrowing season. Richness, evenness, and H diversitywere generally greater on the south plot whereas herbaceous abundance wasgreater on the north plot. Species composition and diversity showed markedphenological (temporal) changes, and comparison of diversity measures at micro-and meso-scales produced markedly different results. Minimum sample areas of150–200 m² were needed to evaluate micro-scalerichness in these species rich communities, suggesting that forest understoriesmay be frequently undersampled in ecological studies. Comparison of observedandestimated meso-scale richness also suggested underestimation of richness in thenorth plot, particularly earlier in the growing season. Thus, sample size,area,and time of sampling appear critical to assessment of diversity in spatiallyandtemporally variable communities such as herbaceous forest understories.     

THE EFFECTS OF LIGHT AND NITROGEN ON GROWTH,PIGMENT CONTENT,AND BIOCHEMICAL COMPOSITION OF GRACILARIA FOLIIFERA V. ANGUSTISSIMA (GIGARTINALES,RHODOPHYTA)1

The combined effects of light intensity and nitrogen (NO₃^?) on growth rate, pigment content, and biochemical composition of Gracilaria foliifera v. angustissima (Harvey) Taylor was investigated using outdoor continuous cultures. Growth of Gracilaria increased linearly with increasing light to 0.43 doublings d^?1 at high light levels (383 ly d^?1 of in situ light), suggesting that light may often limit growth of this plant in nature. Chlorophyll a and phycoerythrin contents were inversely proportional to light level and growth rate. However, pigment content did not affect the growth capacity of Gracilaria. There was no increase in growth or pigment content with increasing additions of nitrogen. The low nitrogen treatment was unenriched seawater that had higher NO₃^? levels than most coastal waters (influent = 8.61 μM; residual = 0.94 μM). When growing near its maximum rate under high light intensities, Gracilaria had a significantly (P < 0.001) lower phycoerythrin: chlorophyll a ratio (phyco: Chl a) than did Gracilaria growing more slowly under lower light (Phyco:Chl a of 2.8 ± 0.2 vs. 3.8 ± 0.3). Faster growing plants also had C:N ratios above 10, indicating N- limitation. In addition to harvesting light the phycobiliproteins of Gracilaria may store nitrogen. Growth rates of Gracilaria correlated negatively with ash (r =–0.85) and positively with the carbon: phycoerythrin ratio (r = 0.85), suggesting that these two indices can be used to estimate growth in the field.