Establishment of native tropical timber trees in monoculture and mixed-species plantations: Small-scale effects on tree performance and insect herbivory

Plantations of native timber species established on former pasture are a potential strategy to reduce the logging pressure on remnant natural forests in the tropics. Such plantations may help to mitigate or reverse the negative impacts of land degradation, and they may contribute to the long-term livelihood of livestock farmers. Planting native trees is, however, perceived as a risky activity due to limited knowledge of their performance and due to marked losses of newly established seedlings attributed to insect pests. Our study focuses on the small-scale effects of environmental heterogeneity, tree diversity and insecticide treatment on the performance of three native Central American timber species two years after establishment, and on damage inflicted by insect herbivores. Growth, survival and herbivore leaf damage were quantified for Anacardium excelsum (Anacardiaceae), Cedrela odorata (Meliaceae) and Tabebuia rosea (Bignoniaceae) planted in an experiment in Panama as (1) monocultures, (2) mixed stands, and (3) mixed stands protected by insecticides. Our study revealed that small-scale effects can have a substantial impact on the success of timber trees planted on former pasture. Growth performance and survival of the three species was strongly affected by small-scale environmental heterogeneity, which was expressed as significant differences in growth and survival among different plantation plots at the same study site. Establishment of trees in mixed stands did not have significant effects on tree survival and growth compared to pure stands, although it reduced herbivore pressure in one of the studied tree species. All tree species grew best and suffered lowest leaf damage when protected by insecticides, indicating a general influence of insect herbivory on growth of newly established trees. In contrast to growth performance, survival was not significantly affected by different management practices or herbivory. The large variability among plots in tree survival and growth, and also in the effects of management practices such as planting design and insecticide treatment, emphasizes the importance of small-scale environmental heterogeneity on tree survival and growth.     

Non-lethal strategies to reduce browse damage in eucalypt plantations

Browsing by mammalian herbivores is a major problem in plantation forestry worldwide. This has traditionally been controlled through a reduction in herbivore numbers achieved by lethal means, namely shooting and poisoning. In recent years, however, there has been increasing social and political pressure to reduce reliance on such lethal controls, and consequently research into non-lethal controls is becoming increasingly important. In order for non-lethal methods to be efficiently incorporated into management, however, we must first identify those that are most effective under operational conditions. Different methods are rarely tested concurrently, and many have only been proven effective with captive animals or in small scale field trials. This study therefore sought to consolidate past research by combining non-lethal methods, which were known to significantly reduce browsing damage, into a single trial to enable their relative effectiveness to be compared under operational conditions.     

Mixed-species plantations of Acacia mangium and Eucalyptus grandis in Brazil: 1. Growth dynamics and aboveground net primary production

Nitrogen fertilizer inputs increased sharply over the last decade in Brazilian eucalypt plantations. Due to the economic and potential environmental cost of fertilizers, mixed plantations with N-fixing species might be an attractive option to improve the long-term soil N status. A randomized block design was set up in southern Brazil, including a replacement series and an additive series design, as well as a nitrogen fertilization treatment. The development of mono-specific stands of Eucalyptus grandis (0A:100E) and Acacia mangium (100A:0E) was compared with mixed plantations in proportions of 1:1 (50A:50E), and other stands with different densities of acacia for the same density of eucalypts. The objective was to assess the effect of inter-specific interactions on the early development of the two species. Aboveground biomass was measured 6, 12, 18 and 30 months after planting, sampling 6–10 trees of each species per treatment at each age, and allometric equations were established in 0A:100E, 100A:0E, 50A:50E and 50A:100E. The height and basal area of E. grandis seedlings were enhanced by 12% and 30%, respectively by N fertilization at age 1 year. Inter-specific competition led to a stratified canopy, with suppression in acacia growth earlier for basal area than for height. The mean number of stems per acacia tree at 36 months after planting was significantly higher in pure stands (3.7), than in 50A:50E (2.7) and in the additive series (between 1.6 and 1.8). H/D ratios were highly sensitive to inter-tree competition for the two species. The suppressed acacia understorey in mixed-species stands did not influence biomass production and partitioning within eucalypts. This pattern led to biomass accumulation combining the two species in 50A:100E that was about 10% higher than in 0A:100E, from age 12 months onwards. Aboveground net primary production (ANPP) amounted to 25 Mg ha⁻¹ and 37 Mg ha⁻¹ from age 18 to 30 months in 100A:0E and 0A:100E, respectively. Acacia ANPP in 50A:100E amounted to 2 Mg ha⁻¹ over the same period, as a result of substantial inter-specific competition. An increment in biomass production in these very fast-growing eucalypt plantations was achieved introducing acacia as an understorey and not in the 50A:50E design, as observed in other studies.     

桉树人工林生长模型研究进展

桉树人工林生长模型包括传统统计模型和现代动态模型。传统统计模型方法包括单因素回归模型、多元回归模型、灰色系统模型、单一分布拟合法、多模型优选法、逐步回归法及其他改进方法。这些统计模型虽然简单、拟合精度高,但只是静态的经验或半经验模型,并未阐明机理,普适性很差,对于动态反馈也无能为力。现代动态模型主要包括3-PG过程模型、人工神经网络模型和系统动力学模型。与传统统计模型相比,动态模型相对较复杂、精度较低,但能够满足多因素交互、非线性预测、动态模拟、多目标决策的研究要求。文中分析了桉树人工林生长模型的特点、各研究方法的比较及系统动力学在林业中的应用,认为系统动力学模型更适合涉及多因素反馈的桉树人工林生长模型的研究。     

A positive growth response to NaCl applications in Eucalyptus plantations established on K-deficient soils

Contrasting responses of Eucalyptus trees to K fertilizer applications have been reported on soils with low K contents. A complete randomized block experiment was set up in Brazil to test the hypothesis that large atmospheric deposits of NaCl in coastal regions might lead to a partial substitution of K by Na in Eucalyptus physiology and enhance tree growth. Treatments with application of 1.5, 3.0, 4.5 kmol K ha⁻¹ (K_1.5, K_3.0, K_4.5, respectively) as KCl, 3.0 kmol K ha⁻¹ applied as K₂SO₄, 3.0 kmol Na ha⁻¹ (Na_3.0) as NaCl commercialized for cattle feeding, and a mixture of 1.5 kmol K + 1.5 kmol Na ha⁻¹ (K_1.5 + Na_1.5) were compared to a control treatment (C) with no K and Na applications. All the plots were fertilized with large amounts of the other nutrients.     

Stand level estimation of root respiration for two subtropical plantations based on in situ measurement of specific root respiration

In this study, the stand level root respiration was estimated for two monoculture plantations: Acacia crassicarpa and Eucalyptus urophylla, based on in situ measurement of specific root respiration using simplified root chamber method. The respiration rates of fine roots (<5 mm) were significantly higher than those of coarse roots (>5 mm) for both A. crassicarpa and E. urophylla species. The root respiration of A. crassicarpa showed a clear seasonal pattern with a higher value in the wet season. For E. urophylla, the seasonal pattern was observed for fine roots but not for coarse roots. After determining the biomass of fine roots and coarse roots and their specific rates of respiration at different time points, root respiration at the stand level (R_a) was estimated using a direct up-scaling model. We found that the R_a accounted for 14% and 19% of total soil respiration (R_s) for A. crassicarpa and E. urophylla, respectively. The fine (R_Tf) and coarse (R_Tc) root respiration at the stand level accounted for about 47% and 53% of the R_a for A. crassicarpa, and accounted for 58% and 42% for E. urophylla. This suggests that coarse root respiration cannot be ignored when estimating the root respiration at the stand level. Our results showed that the Q₁₀ values were more accurate in representing the temperature dependence when the confounding effect of soil moisture was considered. This study introduces an alternative approach to estimate stand level root respiration, but its reliability is largely dependent on the accuracy of root biomass quantification.     

Effects of non-native spruce plantations on small mammal communities in subarctic birch forests

Voles and shrews are key species in northern forest ecosystems. Thus, it is important to quantify to what extent new forestry practices such as planting of non-native tree species impact these small mammals. In northern Norway stands of coastal subarctic birch forests have increasingly been converted to non-native spruce stands during the last century. This leads to changes in the forest floor vegetation and soil conditions that can be expected to negatively impact the community of ground-dwelling small mammals. In this 10-year trapping study we contrasted seasonal small mammal population abundances in spruce plantations with four birch forest varieties. Six different small mammal species were trapped (in descending order of abundance; common shrew Sorex araneus, red vole Myodes rutilus, field vole Microtus agrestis, grey-sided vole M. rufocanus, pygmy shrew S. minutus and water shrew Neomys fodiens). None of the voles appeared to exhibit temporal dynamics resembling population cycles. The three most numerous species were clearly less abundant in the spruce plantations compared to the other forest types. Autumn abundances were most impacted by spruce plantations, indicating that growth rates in the reproductive season were more influenced than winter declines. Species associated with productive forest habitats (i.e. field vole and common shrew) were most impacted by tree species conversion. Still young spruce plantations inter-mixed with birch trees and the ecotone habitat, sustained small mammal abundances comparable to the native birch forests. This implies that managing spruce plantations to maintain a mix of different tree species and high spatial heterogeneity (i.e. more ecotones), will reduce the negative impacts on the small mammal community. On the contrary, if young spruce plantations, as they age become spruce monocultures covering larger parts of the landscapes than they do presently, the negative effects on small mammal communities may be larger than observed in the present study.     

4个桉树无性系愈伤诱导和分化

桉树(Eucalyptus L’Hérit.)是世界三大树种之一,随着生物技术的发展,桉树转基因研究正以前所未有的速度发展,而不同的种、无性系胚性愈伤诱导的难易和植株再生能力的强弱直接影响转基因工程实施的进程。尾巨桉(E.urophylla S.T.Blake×E.grandis W.Hill ex Maiden)无性系DH3229、巨尾桉     

桉树扦插生根和生长性状的QTL定位

本研究以1个尾叶桉×细叶桉F1谱系为材料,在亲本遗传图谱上对扦插穗条的根数、最长根长、生根率和根干质量4个扦插生根性状及13、18、46个月生的树高和胸径2个生长性状的数量性状位点(QTL)进行了定位,结果表明:在母本尾叶桉和父本细叶桉上分别检测到6个和7个扦插生根相关的QTL,似然函数比值的对数值(LOD)2.0～3...     

2种桉树不同林龄生物量与能量的研究

对广东廉江市石岭林场2种桉树不同林龄的生物量和能量进行了研究。结果表明:2种桉树各器官生物量及林分生物量都随林龄的增大而增大,各器官生物量都以干的最大,叶的最小。4.5年生雷林1号桉和尾巨桉林分生物量分别为77.13、80.03 t·hm^-2。不同器官间灰分含量、干质量热值、去灰分热值差异很大,排序分别为:皮>叶>根>枝>干,叶>枝或干>根>皮,叶>枝或根>干>皮。叶、枝、根、干的灰分含量随林龄的增大有减小的趋势,而干质量热值则相反;皮的灰分含量和干质量热值随林龄的变化与前者相反。植     

Predicting growth and sequestration of carbon by plantations growing in regions of low-rainfall in southern Australia

In regions of Australia of low–medium rainfall (500–800 mm/year), there is growing community and land-owner support for re-planting trees to achieve multiple environmental objectives, particularly amelioration of soil salinity. Sequestration of carbon by newly established trees is not only another important environmental benefit, but also a potential commercial benefit. To obtain estimates of carbon sequestered by species of commercial potential in such regions, we calibrated the carbon (C) accounting model FullCAM to Eucalyptus cladocalyx and Corymbia maculata plantations. This was achieved by harvesting trees of a range in sizes to determine the allometric relationships that most accurately predict biomass and stem density from measures of stem diameter. Predictions of stem diameter were obtained from a forest growth model (3-PG) previously calibrated for these two species. By applying these predictions of changes in stem diameter as the stand matures in our allometric relationships, we estimated changes in partitioning of biomass (between stem, branches, bark, foliage and roots) and stem wood density as the stand matures under scenarios of 500, 600 and 750 mm mean annual rainfall. We found that for both species, regardless of annual rainfall, throughout the rotation 37–50% of carbon sequestered in the total tree biomass was in the stem, 18–27% in both branches and roots, and the remainder in foliage or bark. However, rate of accumulation of carbon was dependent on annual rainfall, with average annual rate of sequestration of carbon in tree biomass and litter during the first rotation of E. cladocalyx (or C. maculata) increasing from 3.68 (or 4.17) to 4.72 (or 4.86) Mg C ha⁻¹ yr⁻¹ as annual rainfall increased from about 500 to 750 mm. Although it was predicted that decomposition negated any accumulation of debris between successive rotations, carbon was predicted to accumulate in sawlog products, given that assumed rates of product decomposition were slightly less than their rate of accumulation. This resulted in a slight increase (<8 Mg C ha⁻¹) in predicted total sequestration of carbon between successive rotations.     

Can Eucalyptus invade native forest fragments close to commercial stands?

Some Eucalyptus species are widely used as a plantation crop in tropical and subtropical regions. One reason for this is the diversity of end uses, but the main reason is the high level of wood production obtained from commercial plantings. With the advancement of biotechnology it will be possible to expand the geographical area in which eucalypts can be used as commercial plantation crops, especially in regions with current climatic restrictions. Despite the popularity of eucalypts and their increasing range, questions still exist, in both traditional planting areas and in the new regions: Can eucalypts invade areas of native vegetation, causing damage to natural ecosystems biodiversity?The objective of this study it was to assess whether eucalypts can invade native vegetation fragments in proximity to commercial stands, and what factors promote this invasive growth. Thus, three experiments were established in forest fragments located in three different regions of Brazil. Each experiment was composed of 40 plots (1 m² each one), 20 plots located at the border between the forest fragment and eucalypts plantation, and 20 plots in the interior of the forest fragments. In each experimental site, the plots were paired by two soil exposure conditions, 10 plots in natural conditions and 10 plots with soil exposure (no plant and no litter). During the rainy season, 2 g of eucalypts seeds were sown in each plot, including Eucalyptus grandis or a hybrid of E. urophylla × E. grandis, the most common commercial eucalypt species planted in the three region. At 15, 30, 45, 90, 180, 270 and 360 days after sowing, we assessed the number of seedlings of eucalypts and the number of seedlings of native species resulting from natural regeneration. Fifteen days after sowing, the greatest number of eucalypts seedlings (37 m⁻²) was observed in the plots with lower luminosity and exposed soil. Also, for native species, it was observed that exposed soil improved natural germination reaching the highest number of 163 seedlings per square meter. Site and soil exposure were the factors that have the greatest influence on seed germination of both eucalypt and native species. However, 270 days after sowing, eucalypt seedlings were not observed at any of the three experimental sites. The result shows the inability of eucalypts to adapt to condition outside of their natural range. However, native species demonstrated their strong capacity for natural regeneration in forest fragments under the same conditions where eucalypts were seeded.     

Growth and physiological response of six Australian rainforest tree species to a light gradient

An understanding of growth and photosynthetic potential of subtropical rainforest species to variations in light environment can be useful for determining the sequence of species introductions in rainforest restoration projects and mixed species plantations. We examined the growth and physiology of six Australian subtropical rainforest tree species in a greenhouse consisting of three artificial light environments (10%, 30%, and 60% full sunlight). Morphological responses followed the typical sun-shade dichotomy, with early and late secondary species (Elaeocarpus grandis, Flindersia brayleyana, Flindersia schottiana, and Gmelina leichhardtii) displaying higher relative growth rate (RGR) compared to mature stage species (Cryptocarya erythroxylon and Heritiera trifoliolatum). Growth and photosynthetic performance of most species reached a maximum in 30–60% full sunlight. Physiological responses provided limited evidence of a distinct dichotomy between early and late successional species. E. grandis and F. brayleyana, provided a clear representation of early successional species, with marked increase in A_max in high light and an ability to down regulate photosynthetic machinery in low light conditions. The remaining species (F. schottiana, G. leichhardtii, and H. trifoliolatum) were better represented as falling along a shade-tolerant continuum, with limited ability to adjust physiologically to an increase or decrease in light, maintaining similar A_max across all light environments. Results show that most species belong to a shade-tolerant constituency, with an ability to grow and persist across a wide range of light environments. The species offer a wide range of potential planting scenarios and silvicultural options, with ample potential to achieve rapid canopy closure and rainforest restoration goals.     

Optimizing the joint production of timber and bilberries

Berries and mushrooms are increasingly appreciated products of Finnish forests. Therefore, there is a need to integrate them in silvicultural planning. Bilberry (Vaccinium myrtillus L.) is an economically important wild berry that is widely collected for household consumption and sale in North Karelia, Finland. In this study, bilberry yield models developed recently were included in a stand growth simulator and the joint production of timber and bilberry was optimized by maximizing soil expectation value (SEV) with 3% discounting rate, assuming that 75% of the bilberry yield is harvested. The effect of bilberry production on the optimal stand management increased with increasing bilberry price. With high bilberry prices (4–8 € kg⁻¹) it was optimal to manage the mixed stand of Scots pine, Norway spruce and birch, and the pure stand of Norway spruce so as to promote bilberry production. In the Scots pine stand, where bilberry yields are higher, bilberry production affected optimal stand management already with a price of 2 € kg⁻¹. Compared to timber production, joint production led to longer rotation lengths, higher thinning intensities, more frequent thinnings, and higher share of Scots pine in the mixed stand. The contribution of bilberries to the total SEV increased with increasing bilberry price and discounting rate. In the mixed stand and pine stand the SEV of bilberry production, calculated with 3% discounting rate, exceeded the SEV of timber production when bilberry price was 4 € kg⁻¹.With 4% discounting rate this happened already with bilberry price of 2 € kg⁻¹. It was concluded that forest management which promotes bilberry yields is the most profitable in pine stands where the potential bilberry yields are high.     

Relationships between nursery practices and field performance for Eucalyptus plantations in Brazil

The 3.5 million ha of Eucalyptus plantations in Brazil support many domestic and industrial activities. In the pulpwood sector, forest productivity has increased from 12 to 40 m³ ha^–1 year^–1 through breeding and silviculturalpractices. Nurseries have fostered thisevolution with changes in container/substratetypes, propagation methods and specificinfrastructure for each plant growth phase.Eucalyptus plants are mainly produced bycuttings, using rigid tube containers filledwith vermiculite-organic compound substrates.Stock quality is based on morphologicalcharacteristics of the plants, andenvironmental conditions are highly significantfor plant performance. Performance is dependenton physiological short-term effects related tosurvival and long-term effects of genotype/rootmorphology related to tree development. Currentsilvicultural trends are reducing establishmentcosts but increasing the risk of using lowquality plants. The development of fieldperformance models and nursery physiologicaltests is recommended for Eucalyptusplantations in Brazil. Improved experimentaldesigns are needed to capture the interactionbetween plant quality and field performance.     

Status of floor vegetation under some monoculture and mixculture plantations in North India

A study conducted to assess the floor vegetation shows the presence of 17 species underPopulus deltoides in comparison to 28, 29 and 35 underAlbizia lebbeck, Dalbergia sissoo and mix-culture plantations, respectively. Further, the biomass and density of floor vegetation is considerably reduced underP. deltoides compared to other plantations. In addition, the indices for species richness, evenness, dominance and diversity are also reduced underP. deltoides showing some inhibitory mechanism being operative under these plantations. Based on available literature, allelopathy is speculated to be responsible for this reduction.     

Ungulate herbivory of regenerating conifers in relation to foliar nutrition and terpenoid production

Ungulate browsing greatly influences regeneration dynamics of some forest ecosystems, yet the relationship between browse susceptibility and foliar chemistry of forest tree seedlings is not well understood. We applied field fertilization (15N-9P-10K controlled-release fertilizer at 0, 20, 40, and 60 g per seedling) and investigated how subsequent changes in terpenoid production and foliar nutrition influence ungulate browse preference for Douglas-fir (Pseudotsugamenziesii (Mirb.) Franco), western hemlock (Tsuga heterophylla Raf. Sarg.), and western redcedar (Thujaplicata Donn ex D. Don) seedlings across four sites in northwestern Oregon, USA. Fertilization increased foliar N concentration of all three species, but above-ground growth of only Douglas-fir and western hemlock. Foliar monoterpene concentrations for western hemlock and western redcedar also increased at higher fertilization rates, while Douglas-fir monoterpene production was not affected by fertilization. Regardless of monoterpene levels, ungulate browse preference was greater for fertilized western hemlock seedlings. The opposite response, however, was observed for western redcedar at two of the four study sites where the likelihood of browse was greater for non-fertilized than fertilized seedlings. Differences in browse preference among species may depend on the type and amount of individual monoterpenes manufactured in response to fertilization. Western redcedar produce α- and β-thujone, oxygenated monoterpenes known to promote gastroenteritis and possibly inhibit microbial rumen activity, which were absent from Douglas-fir or western hemlock. Higher concentrations of α- and β-thujone associated with increasing fertilizer rate provide a plausible explanation as to why ungulates preferred non-fertilized western redcedar. Our results illustrate species-specific adaptation in browse avoidance and selective ungulate browsing behavior of individual trees as linked to foliar chemistry.     

Opportunity for hybridization between two oak species in mixed stands as monitored by the timing and intensity of pollen production

The opportunity of cross-pollination in mixed stands of two oak species (cork oak and holm oak) was studied by characterizing individual phenologies of flowering. In the spring of 1998 at one stand consisting of 64 marked trees, there was a period of 19 days when maximal pollen release in one species and stigma receptivity in the other occurred simultaneously, enabling interspecific gene flow in either direction. This happened in spite of an average time separation of 22 days between the two species, reflecting a considerable intraspecific variation in the timings of flowering. Flowering intensities (as estimated from male flower abundance) were high, but fruiting intensities were comparatively low. Shortly after pollination, considerable abortion of female flowers and early fruits was recorded. In 2000, the interspecific overlap of phenologies was drastically reduced due to a delay in cork oak flowering. On the other hand, the individual timings were repeatable for most trees, at least in holm oak. Two other mixed stands were subject of parallel studies, with similar results in all traits except for a less dramatic reduction in fruiting intensities. In spite of the high opportunity for cross-pollination in 1998, and given the lack of hybrids among the progenies from the subsequent fruiting season [Oliveira, P., Custódio, A.C., Branco, C., Reforço, I., Rodrigues, F., Varela, M.C., Meierrose, C., 2003. Hybrids between cork oak and holm oak: isoenzyme analysis. Forest Genet. 10, 283–298], it can be concluded that the prerequisite of cross-pollination is clearly insufficient for hybridization to succeed. Post-pollination processes must play an important role in the maintenance of reproductive isolation between the two species.     

Relating MODIS vegetation index time-series with structure,light absorption and stem production of fast-growing Eucalyptus plantations

By allowing the estimation of forest structural and biophysical characteristics at different temporal and spatial scales, remote sensing may contribute to our understanding and monitoring of planted forests. Here, we studied 9-year time-series of the Normalized Difference Vegetation Index (NDVI) from the Moderate Resolution Imaging Spectroradiometer (MODIS) on a network of 16 stands in fast-growing Eucalyptus plantations in São Paulo State, Brazil. We aimed to examine the relationships between NDVI time-series spanning entire rotations and stand structural characteristics (volume, dominant height, mean annual increment) in these simple forest ecosystems. Our second objective was to examine spatial and temporal variations of light use efficiency for wood production, by comparing time-series of Absorbed Photosynthetically Active Radiation (APAR) with inventory data.     

Why do Pinus species have different growth dominance patterns than Eucalyptus species? A hypothesis based on differential physiological plasticity

It has been observed that Eucalyptus stands show high growth dominance levels while Pinus stands show null or low growth dominance levels. We hypothesized that this differential behaviour is linked to a higher degree of physiological-biochemical plasticity in Eucalyptus than in Pinus species related to photosynthetic capacity. This leads to an increment in growth efficiency (GE) difference between the largest and the smallest trees of a stand, and therefore to high growth dominance levels in Eucalyptus. To test our hypothesis we carried out a bibliographical survey and reanalyzed data from Pinus ponderosa and Pinus taeda plantations in Argentina. We found that some species within the genus Eucalyptus present higher growth dominance levels, physiological plasticity and GE differentiation than Pinus species. The mean maximum values of these traits reported for any Eucalyptus species were: growth dominance coefficient, 0.48; photosynthetic capacity increment when resource availability increases, 175%; GE difference between the largest and the smallest trees of a stand, 300%. Mean maximum values for the same traits reported for any Pinus species were 0.13, no phostosynthetic plasticity as the most frequent pattern, and 51%, respectively. In Pinus species the most frequent response to an increase in resource availability is characterized by an increase in leaf area or biomass, maintaining a similar photosynthetic capacity per unit area. However, it appears that in P. ponderosa there are some situations, characterized by a high degree of intraspecific competition, leading to a very high degree of GE differentiation which deserve future research. Although we did not find any study reporting simultaneously all variables concerning our hypothesis (growth dominance, growth efficiency differentiation and physiological plasticity) any of the circumstantial evidence found in the bibliography contrasts our hypothesis.