We present a meta-analysis of plant responses to fertilization experiments conducted in lowland, species-rich, tropical forests. We also update a key result and present the first species-level ...analyses of tree growth rates for a 15-yr factorial nitrogen (N), phosphorus (P), and potassium (K) experiment conducted in central Panama. The update concerns community-level tree growth rates, which responded significantly to the addition of N and K together after 10 yr of fertilization but not after 15 yr. Our experimental soils are infertile for the region, and species whose regional distributions are strongly associated with low soil P availability dominate the local tree flora. Under these circumstances, we expect muted responses to fertilization, and we predicted species associated with low-P soils would respond most slowly. The data did not support this prediction, species-level tree growth responses to P addition were unrelated to species-level soil P associations. The meta-analysis demonstrated that nutrient limitation is widespread in lowland tropical forests and evaluated two directional hypotheses concerning plant responses to N addition and to P addition. The meta-analysis supported the hypothesis that tree (or biomass) growth rate responses to fertilization are weaker in old growth forests and stronger in secondary forests, where rapid biomass accumulation provides a nutrient sink. The meta-analysis found no support for the long-standing hypothesis that plant responses are stronger for P addition and weaker for N addition. We do not advocate discarding the latter hypothesis. There are only 14 fertilization experiments from lowland, species-rich, tropical forests, 13 of the 14 experiments added nutrients for five or fewer years, and responses vary widely among experiments. Potential fertilization responses should be muted when the species present are well adapted to nutrient-poor soils, as is the case in our experiment, and when pest pressure increases with fertilization, as it does in our experiment. The statistical power and especially the duration of fertilization experiments conducted in old growth, tropical forests might be insufficient to detect the slow, modest growth responses that are to be expected.
Relative growth rates (RGR), i.e., growth rates per unit biomass (ΔM/M, where M is plant mass and ΔM is the change in M over a period of time), reflect growth strategies of plant species. ...Partitioning of RGR to net assimilation rate (ΔM/leaf area) and leaf area ratio (leaf area/M) provides further insights into allocation strategies. To apply this analytical approach to large canopy tree species, we used crown area (Ac) as a proxy for leaf area to understand variations of RGR partitioned to space use efficiency (SUE, ΔM/Ac) and space occupation efficiency (SOE, Ac/M). With UAV imagery, we measured Ac of 226 co-occurring individuals of 14 canopy tree species in a 1-ha stand in a temperate old-growth mixed-forest in Japan, and analyzed how RGR was related to SUE and SOE. The results show that deciduous species exhibited higher SOE and lower SUE compared to evergreen species, even though their RGR values largely overlapped. Late successional species tended to have higher RGR through higher SUE than early-to-middle successional species. We also analyzed the relationship of absolute growth rates (AGR) with several functional traits including DBH (diameter at breast height), Ac, leaf- and stem traits. Both Ac and DBH were strong determinants of AGR across species. Low specific leaf area (SLA, leaf area per unit leaf mass) and high wood density positively contributed to AGR across species, offering long-term growth advantages for old-growth natural forest. The analytical framework introduced in this study may be useful to elucidate the variation of tree growth strategies of canopy trees in natural forest stands.
•Relative growth rates of canopy trees are decomposed into space occupation and use.•Variations in growth strategy are underpinned by functional types and succession.•Late-successional species show high space use than early-mid successional species.•Leaf level area-mass ratio is coupled with crown level area-mass ratio.
Carbonates are only a minor sink of oceanic lithium, yet the presence of this element and its abundance relative to other metal cations in natural carbonate minerals is routinely used as a ...paleo-environmental proxy. To date, however, experimental studies on the influence of physicochemical parameters that may control lithium incorporation in calcite, like pH and precipitation rate, are scarce. Therefore, we experimentally studied Li incorporation in calcite to quantify the apparent partitioning coefficient (DLi∗=cLi/cCacalcitemLi+/mCa2+solution) between calcite and reactive fluid as a function of calcite growth rate and pH. The obtained results suggest that DLi∗ increases with calcite growth rate, according to the expression:LogDLi∗=1.331(±0.116)×LogRate+6.371(±0.880)(R2=0.87;10-8.1≤Rate≤10-7.1molm-2s-1)
Additionally the experimental results suggest that DLi∗ values exhibit a strong pH dependence. For experiments conducted at similar growth rates (i.e. Rate = 10−7.7±0.2 mol m−2 s−1), DLi∗ decreases with increasing pH as described by:LogDLi∗=-0.57(±0.047)×pH+0.759(±0.366)(R2=0.90;6.3<pH<9.5)
The positive correlation of DLi∗ with calcite growth rate is consistent with an increasing entrapment of traces/impurities at rapidly growing calcite surfaces, although the incorporation of monovalent cations such as Li+ and Na+ does not necessarily imply a substitution of Ca2+ ions in the calcite crystal lattice. The dependence of DLi∗ on pH can be considered as an indication that activity of aqueous HCO3− controls the incorporation of Li+ in calcite. The proposed coupled reaction can be explained by charge balance of these monovalent species, which is likely valid at least during the initial step of adsorption on the crystal surface. These new findings shed light on the mechanisms controlling Li incorporation in calcite and have direct implications on the use of Li partitioning coefficients in natural carbonates as an environmental proxy.
The incorporation of magnesium and sulfate in calcite is frequently used to characterize and trace the environmental conditions occurring during carbonate mineral formation. Although both ions are ...simultaneously incorporated in the growing calcite, the effect of sulfate on magnesium incorporation in calcite is still under-explored. In this study, we examine the Mg incorporation in low-Mg calcite as a function of growth rate at 25 °C and 1 bar pCO2 in the presence and absence of aqueous sulfate. The obtained results suggest that high calcite growth rates induce a significant increase in the partitioning coefficient of Mg between the precipitated low-Mg calcite and aqueous solution (i.e. DMg=nMg/nCacalciteMg2+/Ca2+solution). Obtained DMg values exhibit similar dependence to mineral growth rate for experiments performed in the presence and absence of sulfate, however a systematic shift to lower DMg values is observed for calcites formed in the presence of sulfate. The lower DMg values of calcites formed in sulfate-bearing solutions are attributed to the incorporated SO4 ions which provoke expansion in the unit cell along the c-axis of the newly formed calcite. A larger unit cell is unfavorable for the substitution of Ca by smaller Mg ions in calcite. The coupled effect of (i) sulfate uptake during calcite growth and (ii) precipitation rate on DMg can be expressed by the equation DMg=0.03726-0.02345×-logrp-7-XSO4×0.004607+0.002109×-logrp-7; (−8 ≤ log(rp) ≤ −7; 0 ≤ Xso4 ≤ 2.6; T = 25 °C) and is valid for 0.01 ≤ DMg ≤ 0.04. In analogy to Mg also SO4 ions are inhibiting calcite growth, thus promoting the formation of aragonite due to adsorption phenomena and blocking of calcite surface sites. These results improve our understanding on physicochemical parameters controlling CaCO3 composition and mineral polymorphism and are discussed in their relevance for the use of magnesium and sulfate environmental proxies in natural surroundings.
Calcite and aragonite growth rate experiments have been performed in the presence of aqueous Ba at 25 °C using the constant addition technique as a function of CaCO3 mineral growth rate (mol/m2/s). ...The partitioning of Ba in calcite at −8.4 ≤ Logrcalcite ≤ −7.3 exhibits a weak dependence on growth rate that can be expressed as:
LogDBa,calcite=0.2477±0.0543×Logrcalcite-0.2949±0.4222;R2=0.81
In the case of aragonite, Ba partitioning at −9.0 ≤ Lograragonite ≤ −7.8, exhibits a much stronger dependence on growth rate that can be described as:
LogDBa,aragonite=0.4458±0.0563×Lograragonite+3.3407±0.4668;R2=0.84
The determined DBa,aragonite values are systematically lower than unity and come in contrast to previous experimental works where Ba partitioning studied during aragonite nucleation. They are, however, in excellent agreement with the theoretical free energy correlation model by Wang and Xu (2001) that considers the effect of ionic radii size on DBa,aragonite. The defined growth rate dependence of Ba incorporation in aragonite provides new insights on the role of ionic radii size during the incorporation of trace elements and the formation of solid-solutions from aqueous solutions at low oversaturation degrees. The data presented here shed light on the process controlling the elevated DBa,aragonite values occurring at high saturation degrees of natural fluids with respect to aragonite that have been recorded in natural samples. Furthermore, the trend experimentally obtained herein has the potential to record variations in growth rate regimes in natural occurring aragonites and to provide information on the environmental conditions of natural waters in the past.
A rapidly changing climate has the potential to interfere with the timing of environmental cues that ectothermic organisms rely on to initiate and regulate life history events. Short‐lived ectotherms ...that exhibit plasticity in their life history could increase the number of generations per year under warming climate. If many individuals successfully complete an additional generation, the population experiences an additional opportunity to grow, and a warming climate could lead to a demographic bonanza. However, these plastic responses could become maladaptive in temperate regions, where a warmer climate could trigger a developmental pathway that cannot be completed within the growing season, referred to as a developmental trap. Here we incorporated detailed demography into commonly used photothermal models to evaluate these demographic consequences of phenological shifts due to a warming climate on the formerly widespread, multivoltine butterfly (Pieris oleracea). Using species‐specific temperature‐ and photoperiod‐sensitive vital rates, we estimated the number of generations per year and population growth rate over the set of climate conditions experienced during the past 38 years. We predicted that populations in the southern portion of its range have added a fourth generation in recent years, resulting in higher annual population growth rates (demographic bonanzas). We predicted that populations in the Northeast United States have experienced developmental traps, where increases in the thermal window initially caused mortality of the final generation and reduced growth rates. These populations may recover if more growing degree days are added to the year. Our framework for incorporating detailed demography into commonly used photothermal models demonstrates the importance of using both demography and phenology to predict consequences of phenological shifts.
We incorporated demographic components into commonly used phenology models to predict the demographic consequences (i.e. developmental trap vs. demographic bonanza) of phenological shifts in a native multivoltine butterfly under a warming climate. We predicted that populations in the southern limits of its range could become resilient against a warming climate by adding a fourth generation resulting in a demographic bonanza. In the northern ranges, this species may experience diametrically opposed consequences where initial warming would expand the thermal window causing a developmental trap followed by a demographic bonanza if warming continues.
The incorporation of Ni and Co in aragonite was experimentally investigated as a function of mineral growth rate using the constant addition technique at 25 °C and 1 bar pCO2. The distribution ...coefficients of Ni and Co (i.e., DMe=Me/CasolidaMe2+/aCa2+fluid) exhibit a linear correlation with mineral growth rate in the range −9.1 ≤ Log rp ≤ −7.5 mol/m2/s. The obtained results suggest a stronger dependency of DNi to growth rate compared to that observed for DCo. These dependencies can be described as:
Log DNi = 1.247 (±0.152) Log rp + 7.448 (±1.212); R2 = 0.82
Log DCo = 0.312 (±0.047) Log rp + 1.664 (±0.383); R2 = 0.81
The DMe-aragonite values for both Ni and Co are systematically lower than unity and their increase with increasing growth rate is in agreement with the incorporation of elements incompatible with the host mineral structure. Using the dependency of DNi and DCo to the saturation indices (SI) of the reactive solution with respect to aragonite, the DMe under equilibrium were estimated. These estimates yield equilibrium Log DMe values of −3.9 and − 1.0 for Ni and Co, respectively. These experimentally defined Deq values are 1.3 to 4.3 orders of magnitude lower compared to theoretical estimates that were earlier published in the literature. Similar to other incompatible elements, the observed increase of DNi and DCo values with degree of saturation, points towards incorporation related to the density of defect sites on the mineral surface. Finally, the dependencies recorded in this study suggest that DNi and DCo have the potential to be used as a proxy for saturation degree of the reactive solution.
Specialty mushrooms have been implicated in foodborne illness outbreaks in the U.S. in recent years. These mushrooms are available to consumers in both their fresh and dried states. Dehydrating ...mushrooms is a convenient way to increase shelf life. The dehydration process results in a lowered water activity (a w ) of the commodity, creating an environment where both spoilage and pathogenic bacteria cannot proliferate. Prior to food preparation and consumption, these mushrooms are typically rehydrated and possibly stored for later use which could lead to increased levels of pathogens. This study examined the survival and growth of Listeria monocytogenes and Salmonella enterica on dehydrated enoki and wood ear mushrooms during rehydration and subsequent storage. Mushrooms were heat dehydrated, inoculated at 3 log CFU/g, and rehydrated at either 5 or 25°C for 2 h. Rehydrated mushrooms were stored at 5, 10, or 25°C for up to 14 d. L. monocytogenes and S. enterica survived on enoki and wood ear mushroom types during rehydration at 5 and 25°C, with populations often <2.39 log CFU/g. During subsequent storage, no growth was observed on wood ear mushrooms, regardless of the rehydration or storage temperature, with populations remaining <2.39 log CFU/g for both pathogens. When stored at 5°C, no growth was observed for either pathogen on enoki mushrooms. During storage at 10 and 25°C, pathogen growth rates and populations after 14 d were generally significantly higher on the enoki mushrooms rehydrated at 25°C; the highest growth rate (3.56 ± 0.75 log CFU/g/d) and population (9.48 ± 0.62 log CFU/g) after 14 d for either pathogen was observed by S. enterica at 25°C storage temperature. Results indicate a marked difference in pathogen survival and proliferation on the two specialty mushrooms examined in this study and highlight the need for individual product assessments. Data can be used to assist in informing guidelines for time and temperature control for the safety of rehydrated mushrooms.
Magnetosonic (MS) waves with frequencies above the proton gyrofrequency can be locally generated by ring‐beam protons in the Martian heavy ion‐rich ionosphere. In this study, we conduct a parametric ...analysis to investigate the effects of heavy ion concentrations, energy (Erb), and angle (αrb) of ring‐beam protons, and wave normal angle on the excitation features of Martian ionospheric MS waves. We find the growth rates and frequency range of MS waves decrease by including O+ and O2+ ions but are insensitive to their relative concentrations. With increasing Erb or αrb, the growth rates of MS waves show a general dropping tendency. Meanwhile, their frequency and wavenumber range are almost unaffected by increasing Erb but shrink to a narrower range mainly distributed in high frequencies by increasing αrb. Unstable MS waves expand to a wider wavenumber range but shrink to a narrower frequency range as they become more oblique.
Plain Language Summary
Magnetosonic (MS) waves are highly compressible electromagnetic emissions in space. Generally, they are locally excited by ring‐like hot protons in plasmas that lack heavy ions. However, a recent study reveals that they can also be locally excited by ring‐beam protons in the Martian heavy ion‐rich ionosphere. Motivated by these differences, we conduct a detailed parametric analysis to investigate how the heavy ion concentrations, the energy and angle of ring‐beam protons, and the wave normal angle affect the dispersion relation and growth rate of Martian MS wave. We find that O+ and O2+ ions lower the growth rates and frequency range of MS waves but the effects of their relative concentrations are insignificant. In addition to the heavy ions, the excitation features of MS waves strongly depend on the wave normal angle and the distribution properties of ring‐beam protons. These results deepen our understanding of the MS wave excitation features in the unmagnetized planet environment with plentiful heavy ions.
Key Points
O+ and O2+ ions lower the growth rates and frequency range of Magnetosonic (MS) waves but the effects of their relative concentrations are insignificant
The growth rates of MS waves show a general dropping tendency with increasing ring‐beam energy or angle
Unstable MS waves expand to a wider wavenumber range but shrink to a narrower frequency range as they become more oblique
Density-independent and density-dependent population regulation has long been a subject of investigation. We examined density-dependent and density-independent factors on growth rates of pronghorn ( ...Antilocapra americana (Ord, 1815)) using a retrospective analysis of population survey data. Across Idaho, we found that as the proportion of the subpopulation harvested the previous year increased, growth rates increased. Similarly, as fawn recruitment increased, growth rates increased. We also found when the growth rate in the previous year increased, the growth rate the subsequent year decreased. When subpopulations were examined independently, we found that in a low-elevation desert subpopulation, growth rates were influenced by growth the previous year. In an agricultural-dominated site, growth rates were influenced by fawn recruitment in the current year and maximum temperature the previous fall. Growth rates in a mid-elevation shrub–steppe site were influenced by drought severity prior to parturition and the growth rates from the previous year. Growth rates in two mountain valley subpopulations were influenced by measures of vegetative greenness. At the statewide scale, while managers may strive for increased numbers of pronghorn, density dependence will limit the ability for a region-wide numerical response. On the localized scale, drivers of growth were temporally and spatially explicit, and biologists must consider site-specific actions.