The effect of water and nitrogen amendments on photosynthesis,leaf demography,and resource-use efficiency in Larrea tridentata,a desert evergreen shrub

Summary In the Chihuahuan Desert of southern New Mexico, both water and nitrogen limit the primary productivity of Larrea tridentata, a xerophytic evergreen shrub. Net photosynthesis was positively correlated to leaf N, but only in plants that received supplemental water. Nutrient-use efficiency, defined as photosynthetic carbon gain per unit N invested in leaf tissue, declined with increasing leaf N. However, water-use efficiency, defined as the ratio of photosynthesis to transpiration, increased with increasing leaf N, and thus these two measures of resource-use efficiency were inversely correlated. Resorption efficiency was not significantly altered over the nutrient gradient, nor was it affected by irrigation treatments. Leaf longevity decreased significantly with fertilization although the absolute magnitude of this decrease was fairly small, in part due to a large background of insect-induced mortality. Age-specific gas exchange measurements support the hypothesis that leaf aging represents a redistribution of resources, rather than actual deterioration or declining resource-use efficiency.     

Mineral nutrition and leaf longevity in an evergreen shrub,Ledum palustre ssp. decumbens

Summary Effects of variable mineral nutrient status on evergreen leaf longevity were investigated in a field fertilization experiment, and by comparison of plants from several sites. The species studied was Ledum palustre spp. decumbens, with a normal leaf life expectancy of 2.06 years and a maximum leaf longevity of 4.5–5.0 years. Most leaf losses took place during the growing season, not during the winter. Fertilization increased leaf production but decreased leaf survivorship. Total number of leaves per stem was unchanged with fertilization. In a comparison among sites, there was a moderate negative correlation between plant N and P concentrations and leaf longevity. These intraspecific responses are similar to known interspecific changes along nutrient gradients, i.e. with high nutrient availability a vegetation should become more deciduous and less evergreen.     

重庆石灰岩地区主要木本植物叶片性状及养分再吸收特征

以重庆石灰岩地区15种常绿木本植物和14种落叶木本植物为研究对象,对两种生活型植物叶片衰老前后叶干物质含量(LDMC)、比叶面积(SLA)和叶片厚度(LT)进行了比较,并采用不同的计算方法(单位质量叶片养分含量、单位面积叶片养分含量)分析了两类植物叶片衰老前后养分含量及再吸收特征,最后对养分再吸收效率与其他叶性状因子之间的关系进行了相关分析。结果表明:常绿植物成熟叶LDMC、LT及衰老叶LT显著低于落叶植物,落叶植物成熟叶和衰老叶SLA均显著高于常绿植物(P0.05);基于单位质量叶片计算的养分含量,常绿植物成熟和衰老叶N、P量均低于落叶植物,而基于单位面积叶片计算的N、P含量则表现出相反的趋势;基于不同方法计算的N、P再吸收效率差异不明显,其中常绿植物基于单位质量叶片养分含量计算的N、P平均再吸收效率为39.42%、43.79%,落叶植物的为24.08%、33.59%;常绿和落叶植物N、P再吸收效率与LDMC、SLA、LT和成熟叶N、P含量之间没有显著相关性,但与衰老叶养分含量存在显著负相关(P0.05)。研究发现,无论是常绿植物还是落叶植物,衰老叶N、P含量均较低,表明石灰岩地区植物具有较高的养分再吸收程度。     

氮磷施肥对拟南芥叶片碳氮磷化学计量特征的影响

研究植物碳(C)氮(N)磷(P)化学计量特征, 有助于了解C、N、P元素的分配规律和确定限制植物生长的元素类型, 理解生长速率调控的内在机制。该研究基于盆栽施肥试验, 测定不同N、P供应水平下拟南芥(Arabidopsis thaliana)叶片的生物量和C、N、P含量, 分析拟南芥的限制元素类型、验证生长速率假说、探讨N、P的内稳性差异和C、N、P元素间的异速生长关系。主要结果如下: 盆栽试验基质中限制元素是P, 施N过多可能引起毒害作用; 拟南芥的生长符合生长速率假说, 即随着叶片N:P和C:P的增加, 比生长速率显著减小; 叶片P含量存在显著的调整系数(3.5), 但叶片N含量与基质N含量之间无显著相关; 叶片N和P含量具有显著的异速生长关系, 但不符合N-P^3/4关系, 施P肥导致表征N、P异速生长关系的幂指数(0.209)显著低于施N肥处理(0.466)。该研究首次基于温室培养实验分析了拟南芥C、N、P的化学计量特征及其对N、P添加的响应, 研究结果将为野外研究不同物种、群落或生态系统的化学计量特征提供参考。     

Resorption proficiency and efficiency of leaf nutrients in woody plants in eastern China

Aims (i) To explore variations in nutrient resorption of woody plants and their relationship with nutrient limitation and (ii) to identify the factors that control these variations in forests of eastern China.Methods We measured nitrogen (N) and phosphorus (P) concentrations in both green and senesced leaves of 172 woody species at 10 forest sites across eastern China. We compared the nutrient resorption proficiency (NuRP) and efficiency (NuRE) of N and P in plant leaves for different functional groups; we further investigated the latitudinal and altitudinal variations in NuRP and NuRE and the impacts of climate, soil and plant types on leaf nutrient resorptions.Important findings On average, the leaf N resorption proficiency (NRP) and P resorption proficiency (PRP) of woody plants in eastern China were 11.1mg g ? 1 and 0.65 mg g ? 1, respectively; and the corresponding N resorption efficiency (NRE) and P resorption efficiency (PRE) were 49.1% and 51.0%, respectively. Angiosperms have higher NRP (are less proficient) values and lower NRE and PRE values than gymnosperms, but there are no significant differences in NRP, PRP and PRE values between species with different leaf habits (evergreen vs. deciduous angiosperms). Trees have higher NRE and PRE than shrubs. Significant geographical patterns of plant nutrient resorption exist in forests of eastern China. In general, NRP and PRE decrease and PRP and NRE increase with increasing latitude/altitude for all woody species and for the different plant groups. Plant functional groups show more controls than environmental factors (climate and soil) on the N resorption traits (NRP and NRE), while site-related variables present more controls than plant types on PRP and PRE. NRP increases and PRP and NRE decrease significantly with increasing temperature and precipitation for the overall plants and for most groups, except that significant PRE–climate relationship holds for only evergreen angiosperms. Leaf nutrient resorption did not show consistent responses in relation to soil total N and P stoichiometry, probably because the resorption process is regulated by the relative costs of drawing nutrients from soil versus from senescing leaves. These results support our hypothesis that plants growing in P-limited habitats (low latitudes/altitudes or areas with high precipitation/temperature) should have lower PRP and higher PRE, compared with their counterparts in relatively N-limited places (high latitudes/altitudes or areas with low precipitation/temperature). Our findings can improve the understanding of variations in N and P resorption and their responses to global change, and thus facilitate to incorporate these nutrient resorption processes into future biogeochemical models.     

Response of leaf anatomy of <Emphasis Type="Italic">Chenopodium acuminatum</Emphasis> to soil resource availability in a semi-arid grassland

Soil water (W), nitrogen (N), and phosphorus (P) availability were manipulated for 2 years in a factorial experiment in a semi-arid grassland in the southeastern Keerqin Sandy Lands, China, to determine what soil resource(s) limits plant growth. We quantified the responses of leaf chemical and anatomical traits for the dominant annual herb Chenopodium acuminatum in this ecosystem to test hypotheses about leaf form and function in relation to soil resource limitation. Nitrogen was the principal limiting factor, and N treatments (alone and in combination with W and P) elicited consistent increases in leaf size and decreases in leaf thickness and water content. Differentiation in the leaf mesophyll between palisade and spongy parenchyma was lacking in N deficient plants and stimulated in all +N treatments. The greater tissue density and cell packing may promote more efficient light capture with improved N nutrition. Mixed additions of W, N, and P were generally more advantageous to leaf expansion and nutrient balance than their isolated additions, therefore water and nutrients synchronous management would be a better approach to promote plant growth at the semi-arid region.     

Nutrient resorption of two evergreen shrubs in response to long-term fertilization in a bog

Plant resorption of multiple nutrients during leaf senescence has been established but stoichiometric changes among N, P and K during resorption and after fertilization are poorly understood. We anticipated that increased N supply would lead to further P limitation or co-limitation with N or K [i.e. P-(co)limitation], decrease N resorption and increase P and K resorption, while P and K addition would decrease P and K resorption and increase N resorption. Furthermore, Ca would accumulate while Mg would be resorbed during leaf senescence, irrespective of fertilization. We investigated the effect of N, P and K addition on resorption in two evergreen shrubs (Chamaedaphne calyculata and Rhododendron groenlandicum) in a long-term fertilization experiment at Mer Bleue bog, Ontario, Canada. In general, N addition caused further P-(co)limitation, increased P and K resorption efficiency but did not affect N resorption. P and K addition did not shift the system to N limitation and affect K resorption, but reduced P resorption proficiency. C. calyculata resorbed both Ca and Mg while R. groenlandicum resorbed neither. C. calyculata showed a higher resorption than R. groenlandicum, suggesting it is better adapted to nutrient deficiency than R. groenlandicum. Resorption during leaf senescence decreased N:P, N:K and K:P ratios. The limited response of N and K and the response of P resorption to fertilization reflect the stoichiometric coupling of nutrient cycling, which varies among the two shrub species; changes in species composition may affect nutrient cycling in bogs.     

Effects of water and mineral nutrient supply on a deciduous and an evergreen dwarf shrub: Vaccinium uliginosum L. and V. vitisidaea L.

The effects of irrigation and fertilization on nutrient content, shoot growth and photosynthetic rate of the deciduous Vaccinium uliginosum L. and the evergreen V. vitis-idaea L. were studied in a field experiment at Abisko, Swedish Lapland. V. vitis-idaea responded to fertilization with a greater increase in leaf nitrogen, phosphorus and potassium concentrations, as well as with a relatively greater change in shoot growth than V. uliginosum . Also to irrigation did V. vitis-idaea respond more strongly. Decreased leaf longevity was indicated in both irrigated and fertilized plants of V. vitis-idaea . It is concluded that no general response pattern for evergreen and deciduous plants can be expected to emerge unless one also takes other plant characteristics, and also site characteristics, into consideration.     

Time‐dependent effects of fertilization on plant biomass in floating fens

Abstract. A cross‐over fertilization experiment was carried out in Dutch floating fens to investigate effects on biomass production in the same and the following years. In total 16 fertilizer treatments were applied, combining four treatments in 1999 with four treatments in 2000 (addition of 20 g.m^?2 N, 5 g.m^?2 P, both elements and unfertilized control). The above‐ground biomass production of vascular plants was co‐limited by N and P in both years. However, in plots that were only fertilized in 1999 the effects of individual nutrients differed between the two years: N‐fertilization slightly increased the amount of biomass produced in the same year (1999), whereas P‐fertilization did so in the following year (2000). Fertilizer applied in 1999 also influenced the effects of fertilizer applied in 2000. One year after N‐fertilization vascular plant growth was still co‐limited by N and P, but one year after P‐fertilization, vascular plant growth was only limited by N. Bryophyte biomass responded weakly to fertilization. Nutrient concentrations in plant biomass, nutrient standing crops and measurements of N and P availability in the soil indicated that one year after fertilization, the N‐fertilizer had mostly ‘disappeared’ from N‐fertilized plots, whereas the availability of P remained markedly enhanced in P‐fertilized plots. In addition, P‐fertilization enhanced the uptake of N by plants the following year. The time‐dependence of fertilizer effects was probably caused by (1) higher addition of P than of N relative to the requirements of plants; (2) longer retention of P than of N in the system; (3) positive effect of P‐fertilization on the availability of N; (4) contrasting effects of N‐ and P‐fertilization on nutrient losses by plants and/or on their responses to subsequent nutrient addition; (5) changing interactions between vascular plants and mosses (mainly Sphagnum spp.); (6) nutrient export through the repeated harvest of above‐ground biomass. To determine which nutrient limits plant growth fertilization experiments should be short, avoiding that indirect effects of a non‐limiting nutrient influence results. To indicate how changed nutrient supply will affect an ecosystem longer‐term experiments are needed, so that indirect effects have time to develop and be detected.     

Few long-term effects of simulated climate change on volatile organic compound emissions and leaf chemistry of three subarctic dwarf shrubs

Climate change is exposing arctic ecosystems to higher temperature, increased nutrient availability and shading due to the increasing cloud cover and the expanding forests. In this work, we assessed how these factors affect the emissions of biogenic volatile organic compounds (BVOCs) from three subarctic dwarf shrub species in a field experiment after 18 treatment years. Of the studied species the willow Salix phylicifolia L. was the only isoprene-emitter with an emission potential of 16.1 ± 8.4 μg g⁻¹ dw h⁻¹ (at 30 °C and photosynthetic photon flux density of 1000 μmol m⁻² s⁻¹). The dwarf birch Betula nana L. had significant emissions of various reactive BVOCs, including monoterpenes and sesquiterpenes. The evergreen Cassiope tetragona (L.) D. Don emitted high amounts of mono- and sesquiterpenes. Due to chance, the temperature in the warming treatment (employing open-top plastic tents) and the unwarmed treatments was similar at the time of the measurements, and therefore long-term indirect effects of warming could be assessed without interference of temperature differences at the time of measurement. The long-term warming had not altered foliar N, P or condensed tannin concentrations, but it had led to other chemical changes detected in the near-infrared reflectance spectra of the leaves. Nevertheless, there were no significant differences in the BVOC emissions per unit leaf mass measured by the dynamic enclosure method and gas chromatography-mass spectrometry. Annual additions of NPK fertilizer, which mimicked increased nutrient availability, had accumulated P in the leaves of all species. In addition, fertilization marginally increased the leaf N concentration of B. nana. The only significant fertilization effect on BVOC emissions was a stimulation of emission of the sesquiterpene β-selinene from S. phylicifolia. The shading treatment obtained by dome-shaped hessian tents did not cause clear long-term changes in leaf chemistry or BVOC emissions. The only observed change was a marginally significant increase in sesquiterpene emissions from B. nana. When the treatment effects on long-term biomass changes in the different treatments were taken into account by proportioning the BVOC emissions to the biomass of each species in the field treatments, warming led to a significant increase and shading to a decrease in the total BVOC emissions per unit ground area from B. nana. These results highlight the importance of an integrated approach for realistic assessment of responses to climate change.     

Declining soil fertility does not increase leaf lifespan within species: evidence from the Franz Josef chronosequence, New Zealand

Leaf lifespan varies widely among plant species, from a few weeks to >40 years. This variation is associated with differences in plant form and function, and the distribution of species along resource gradients. Longer leaf lifespans increase the residence time of nutrients and are one mechanism by which plants conserve nutrients; consequently, leaf lifespan should increase within species with declining soil nutrient availability. The Franz Josef chronosequence is a series of post-glacial surfaces along which soil fertility declines strongly with increasing soil age. We used this fertility gradient to test whether leaf lifespans of six common indigenous woody species increased as soil nutrient availability declined. Leaf lifespan varied from 12.4 months in Coprosma foetidissima (Rubiaceae) to 47.1 months in Pseudopanax crassifolius (Araliaceae). These leaf lifespans sample 12% of the full range of leaf lifespans reported globally and occupy a relatively conservative portion of global leaf trait space. Contrary to our expectations, leaf lifespan of two species (Pseudopanax crassifolius and Prumnopitys ferruginea) decreased by 44?61% with increasing soil age and there were no other relationships between soil age and leaf lifespan. Across all species, leaf nutrient residence times increased by 85% for N and 90% for P with declining soil fertility, but this was caused by increased nutrient resorption efficiency rather than by increased leaf longevity. These data demonstrate that plants increase leaf nutrient resorption efficiency rather than leaf lifespan as a within-species response to long-term declines in soil fertility.     

常绿阔叶林植物叶片N、P化学计量特征对毛竹扩张的响应

为从生态化学计量内平衡角度解释常绿阔叶林不同层次植物对毛竹(Phyllostachys edulis)扩张的生存响应差异性,该研究采用空间代替时间的方法,在江西井冈山国家级自然保护区沿毛竹扩张方向选取典型毛竹-常绿阔叶林界面,依次设置毛竹林、竹阔混交林和常绿阔叶林样地,比较分析了毛竹扩张方向上样地内不同乔木层、灌木层、草本层植物叶片及土壤N、P含量及比例。结果表明:(1)从毛竹林到阔叶林,土壤N含量上升,P含量下降,N:P上升(P<0.05); 乔木层树种 [红楠(Machilus thunbergii)、赤杨叶(Alniphyllum fortunei)及交让木(Daphniphyllum macropodum)]叶片P含量下降,N:P上升(P<0.05); 除灌木层的红果山胡椒(Lindera erythrocarpa)外,各林分中的灌木层和草本层植物N、P含量及比例变化较小。(2)土壤N:P与乔木层、草本层和灌木层植物叶片N:P分布呈显著正相关、负相关与不相关。(3)在各林分中,毛竹叶片N、P含量及比例较稳定。综上认为,毛竹通过改变土壤N、P化学计量特征进行扩张,引起植物体N、P元素化学计量特征发生变化。灌木及草本植物受土壤异质性影响较小,但是乔木层植物N、P元素化学计量特征却因此失衡,这可能是阔叶林乔木层树种存亡受威胁的重要原因。     

Changes in leaf nutrient traits in a wildfire chronosequence

The effect of wildfire on ecosystem function is gaining interest since climate change is expected to increase fire frequency and intensity in many forest systems. Fire alters the nutritional status of forest ecosystems, affecting ecosystem function and productivity, but further studies evaluating changes in leaf nutrient traits induced by forest wildfires are still needed. We used a 17-year-old Pinus canariensis wildfire chronosequence to elucidate the nature of nutrient limitations in natural and unmanaged pine forest in the Canary Islands. Pine needles were sampled in winter and spring and analysed for N and P concentrations. As expected, we found the lowest leaf N and leaf P in recently burned plots. However, the leaf N:P ratio was higher in burned versus unburned plots, suggesting that the decrease in P availability due to the fire is larger than that of N. For all leaf traits and sampling dates, leaf trait values in burned plots matched those observed in unburned plots 17 years after a fire. The N:P ratio found in P. canariensis needles was one of the lowest values reported in the literature for woody species, and suggests that all pine trees in the chronosequence are unambiguously limited by low N availability. Our results show that these N-limited pine forests retained N more efficiently than P 4 years after a wildfire; however, leaf N recovery is slower than P recovery, suggesting that the mechanisms responsible for pine N limitation operate continuously in these forests.     

Effects of nitrogen and phosphorus supply on growth and flowering phenology of the snowbed forb Gnaphalium supinum L.

The warming-induced increase in nutrient mineralization and the further increase in atmospheric nitrogen depositions raise the topic of whether and how alpine plants will react to enhanced nutrient availability. Despite several studies have shown the effects of fertilization on primary production of alpine plants, only few studies have considered the influences of nutrients on reproduction. Here, we investigated the effects of nitrogen (N) and phosphorus (P) amendments on cover, number of ramets, flowering effort and phenological timing of Gnaphalium supinum, an arctic-alpine widespread snowbed species. We set up an experimental design with four fertilization treatments (low N, P without additional N, low N + P, and high N + P) and an unfertilized control for three years (2003–2005), within a late snowbed located in the Italian Alps (Gavia Pass, 2700 m a.s.l.). The cover of Gnaphalium supinum was recorded at the peak of the aboveground biomass development in the three years, while the temporal dynamic of ramet density and reproductive phenophases were monitored during the 2005 growing season. The clonal growth of G. supinum resulted to be co-limited by N and P, while the flowering effort was stimulated by P. Flowering date was advanced by P supply, while N alone did not show any significant effect on phenology. In a warming scenario, with a predicted increase in N and P availability by nutrient mineralization and atmospheric deposition, this species should probably experience some benefits for its growth and reproduction if not limited by other factors such as the length of the growing season or interspecific competition.     

The Glycine-Glomus-Rhizobium Symbiosis : VII. Photosynthetic Nutrient-Use Efficiency in Nodulated, Mycorrhizal Soybeans

Four consecutive trifoliate leaves of 56-day-old symbiotic or nonsymbiotic soybean plants were evaluated individually for CO₂ exchange rates (CER), leaf area and dry weight, and leaf N, P, and starch concentrations. Plants had been inoculated with the vesicular-arbuscular mycorrhizal (VAM) fungus Glomus mosseae and Rhizobium japonicum, with either of the endophytes alone, or with neither at time of planting. Plants lacking one or both endophytes received N and/or P fertilizers to produce plants of equal total leaf dry weight in all four treatments. Photosynthetic P-use efficiency (CER per unit leaf P) was higher in the leaves of VAM plants than in P-fertilized plants regardless of the N source (N₂ fixation or combined N). Photosynthetic N-use efficiency was also higher in VAM than in non-VAM plants, but it was affected by the N source, with higher CER in the nodulated plants. The greatest differences in CER, starch accumulation and leaf area were found between the nonsymbiotic plants and those with both endophytes. Statistical evaluations of leaf parameters for treatment or nutrient concentration (N and P) effects between the tri-partite and the nonsymbiotic treatments showed significant changes in concentration of P, but not N, with decreasing leaf age. Both endophytes apparently enhance CO₂ fixation at N and/or P concentrations lower than those of the nonsymbiotic plants. The effects of the endophytes on CO₂ fixation were additive.     

Significance of sequential leaf development for nutrient balance of the cotton sedge,Eriophorum vaginatum L.

Summary The sedgeEriophorum vaginatum in an interior Alaskan muskeg site produced leaves sequentially at about 1.5-month intervals. Each leaf remained active for two growing seasons. Young leaves (even those initiated late in the season) always had high concentrations of N, P, K and Mg and were low in Ca. Stems had high concentrations of nutrients, sugar, amino acid N and soluble organic P in autumn and spring but low concentrations in summer. Growth of leaves in spring was strongly supported by translocation from storage. Leaves approached their maximum nutrient pool before nutrient uptake began in late spring, one month before maximum biomass. Retranslocation of nutrients from aging leaves could support nutrient input into new, actively growing leaves as a consequence of the sequential leaf development. For instance retranslocation from aging leaves accounted for more than 90 and 85% of P and N input to new leaves appearing in early summer and 100% to leaves that appeared later. Leaching losses were negligible. Half time for decay of standing dead litter was 10 years. We suggest that sequential leaf development paired with highly efficient remobilization of nutrients from senescing leaves enables plants to recycle nutrients within the shoot and minimize dependence upon soil nutrients. This may be an important mechanism enablingEriophorum vaginatum to dominate nutrient-poor sites. This may also explain why graminoids with sequential leaf production cooccur with evergreen shrubs and dominate over forbs and deciduous shrubs in nutrient-poor sites in the boreal forest (e.g., in bogs) and at the northern limit of the tundra zone.     

天童常绿阔叶林若干树种的叶片营养转移研究

对天童国家森林公园 1 8种植物在落叶前后营养转移量的研究表明 ,不同植物的营养转移率不尽相同 ,N的平均转移率为 3 7.86% ,标准偏差 1 0 .67% ;P的平均转移率为 44.76% ,标准偏差 1 5 .40 %。经方差分析 ,N、P转移率无明显差异 (s=0 .1 3 8)。同时 ,P的转移率与植物成熟叶中的P含量及N/P存在正相关 ,而N的转移率与植物成熟叶子的N含量和N/P不相关。另外 ,常绿植物的N转移率平均值是 3 5 .74% (标准差9.46% ) ,落叶植物N转移率平均值是 3 8.72 % (标准差 1 2 .65 % ) ;常绿植物的P转移率平均值是 3 7.72 % (标准差 1 3 .0 0 % ) ,落叶植物P转移率平均值是 5 5 .3 7% (标准差 1 5 .5 4% )。对落叶和常绿阔叶二种生活型植物进行ANOVA分析 ,表明N转移率无明显差异 ,而P转移率有差异 (P <0 .0 5 )。     

Effects of leaf longevity and retranslocation efficiency on the retention time of nutrients in the leaf biomass of different woody species

Summary A study was made of the retention times of N and P in the leaf biomass and their relationship with the retranslocation percentages and the leaf longevities in some woody species in Central Spain. The retention times of both nutrients were strongly related to the nutrient status of each species. These results suggest that a prolonged retention time is a way of increasing nutrient use efficiency in conditions of low nutrient availability. Plants can increase the retention time of nutrients in their leaf biomass by means of an increase in leaf longevity and/or by means of an increase in retranslocation efficiency. However, the effect of the retranslocation efficiency on retention times was almost negligible compared with the effect of leaf longevity. This suggests that an increase in leaf longevity is probably the best adaptation for increasing efficiency in the use of nutrients.     

Coordination of leaf N,anatomy, photosynthetic capacity,and hydraulics enhances evergreen expansive potential

Key message

Reduced leaf longevity, N-fixation, and enhanced hydraulic capacity combined support greater shifts in seasonal photosynthetic capacity of an expansive understory evergreen woody species relative to co-occurring less expansive evergreen species.

Abstract

Physiological functioning typically declines with increased leaf life span. While an evergreen leaf habit is generally associated with reduced leaf N, physiological capacity, and slower growth, most expansive woody species are evergreens and/or N fixers. An evergreen leaf habit enables year-round activity and less investment in carbon and nutrients, while N-fixation enhances photosynthetic capacity. Our objective was to compare anatomy and physiology of three woody evergreens Ilex opaca Aiton (Aquifoliaceae), Kalmia latifolia L. (Ericaceae), and Myrica cerifera (Myricaceae) of varying leaf longevity, N-fixation capability, and known expansive potential in a deciduous forest understory to determine if seasonal physiological performance integrated these factors. We hypothesized that I. opaca (non-expansive) and K. latifolia (moderately expansive), which have longer leaf longevities, would have reduced physiological performance compared to M. cerifera (expansive), which has shorter leaf longevity, and symbiotically fixes atmospheric N. Stomatal conductance to water vapor, photosynthetic and hydraulic capacities, specific leaf area, and leaf %N decreased with increasing leaf life span; however, trends among species were not consistent seasonally. While hydraulic capacity remained constant throughout the year, photosynthetic capacity did not. During the growing season, M. cerifera displayed photosynthetic capacity similar to deciduous species, yet, during the winter, photosynthetic capacity was similar to the slower-growing evergreens. Reduced leaf life span, enhanced hydraulic capacity, and nitrogen fixation support the seasonal shift in photosynthetic capacity observed in M. cerifera. This “hybrid” strategy enables M. cerifera to maximize productivity during months of optimal conditions, thereby promoting rapid growth and expansion in the understory.