A hierarchical perspective in allocation to reproduction from whole plant to fruit and seed level

Allocation to reproduction is often examined at the whole plant level from an allometric perspective; however, investment in reproduction is a hierarchical process which represents the integration of adjustments at different hierarchical levels. This essay review presents some key issues in reproductive allocation by examining problems of allocation at different hierarchical levels within plant. At the whole plant level, the application of ‘Rensch's rule’ to plants is discussed as a mechanism that might explain size differences in relation to gender in dioecious species. At branch level, the existence of vegetative branches may be explained by hypotheses on branch specialization and costs of reproduction. However, vegetative branches might also constitute an ‘assurance’ for reproduction in variable environments. At a lower hierarchical level, fruit abortion is a process widely observed in plants that impose limits to our capacity to estimate and develop general laws about reproductive allocation in plants. The question of whether fruit abortion can be used by the parent to produce a more homogeneous progeny that limits asymmetric competition among sibs is discussed. Finally, it seems that in some cases the allocation within seeds is in accordance with some predictions of kin selection theory. Most of these topics contain open questions that deserve further research.     

植物的表型可塑性、异速生长及其入侵能力

表型可塑性是指同一个基因型对不同环境响应产生不同表型的特性,特定性状的可塑性本身可以遗传,也可以接受选择而发生进化。植物个体的异速生长是指生物体某一特征的相对生长速率不等于第二种特征的相对生长速率的特性,该特性是由物种的遗传性决定的一种固定特征,植物往往朝着最佳的异速生长曲线进化。植物特定基因型在不同环境下,诸如生物量分配和种群几何学上的一些表型差异,既可由异速生长造成,也可由表型可塑性造成。植物本身的异速生长是一种"外观可塑性",而异速生长曲线的改变才是真正的可塑性。植物的表型可塑性、异速生长对于入侵植物的适应具有重要意义。干扰等异质性生境下表型可塑性成为物种生存扩散的有利性状,表型可塑性强的物种更有可能成为广布种。植物本身的异速生长特性或其异速生长曲线的改变都能影响其入侵能力。     

Plasticity and ontogenetic drift of biomass allocation in response to above- and below-ground resource availabilities in perennial herbs: a case study of <Emphasis Type="Italic">Alternanthera philoxeroides</Emphasis>

Changes in plant biomass allocation in response to varying resource availabilities may result from ontogenetic drift caused by allometric growth (i.e., apparent plasticity), a true adjustment of ontogenetic trajectories (true plasticity) or both (complex plasticity). Given that the root allocation of annual species usually decreases during the growth, the developmentally explicit model predicts that annual herbs will exhibit true plasticity in root allocation under above-ground resource limitation and apparent plasticity for moderate stress of below-ground resource. For perennial species, the root allocation of which increases during growth, the reverse patterns would be expected. In this study, we tested the developmentally explicit model with a perennial weed, Alternanthera philoxeroides (Mart.) Griseb. We report its adaptive changes and ontogenetic drift of root allocation in response to different resource levels (i.e., light, water and nutrient availability) by comparing root allocation on both an age and a size basis. The root allocation of A. philoxeroides increased with the size (i.e., ontogenetic drift) during the growth, and exhibited significant changes in response to different resource availabilities. Furthermore, the root allocation in response to water or nutrient availability exhibited typical complex plasticity, while the light stress only slowed down the growth, with the ontogenetic trajectory unchanged (apparent plasticity). The contrasting responses to above-ground and below-ground stresses were consistent with the prediction of the developmentally explicit model.     

Analysing the allometry of multiple interacting traits

Since form and function are tightly integrated in plants, and since plant attributes often scale allometrically, it follows that plant allometry is inherently multivariate. Unfortunately, traditional statistical methods for studying allometric relationships are very restrictive and do not allow one to model multivariate allometric patterns that follow realistic biological hypotheses. In this paper I describe a new statistical test (‘d-sep test’) that allows one to test, and potentially falsify, alternative multivariate orderings of cause-and-effect in the context of allometry.     

Arbuscular mycorrhizal fungi alter plant allometry and biomass-density relationships

Background and Aims

Plant biomass–density relationships during self-thinning are determined mainly by allometry. Both allometry and biomass–density relationship have been shown to vary with abiotic conditions, but the effects of biotic interactions have not been investigated. Arbuscular mycorrhizal fungi (AMF) can promote plant growth and affect plant form. Here experiments were carried out to test whether AMF affect plant allometry and the self-thinning trajectory.

Methods

Two experiments were conducted on Medicago sativa L., a leguminous species known to be highly dependent on mycorrhiza. Two mycorrhizal levels were obtained by applying benomyl (low AMF) or not (high AMF). Experiment 1 investigated the effects of AMF on plant growth in the absence of competition. Experiment 2 was a factorial design with two mycorrhizal levels and two plant densities (6000 and 17 500 seeds m⁻²). Shoot biomass, root biomass and canopy radius were measured 30, 60, 90 and 120 d after sowing. The allometric relationships among these aspects of size were estimated by standardized major axis regression on log-transformed data.

Key Results

Shoot biomass in the absence of competition was lower under low AMF treatment. In self-thinning populations, the slope of the log (mean shoot biomass) vs. log density relationship was significantly steeper for the high AMF treatment (slope = –1·480) than for the low AMF treatment (–1·133). The canopy radius–biomass allometric exponents were not significantly affected by AMF level, but the root–shoot allometric exponent was higher in the low AMF treatment. With a high level of AMF, the biomass–density exponent can be predicted from the above-ground allometric model of self-thinning, while this was not the case when AMF were reduced by fungicide.

Conclusions

AMF affected the importance of below-ground relative to above-ground interactions and changed root vs. shoot allocation. This changed allometric allocation of biomass and altered the self-thinning trajectory.     

The effect of nutrient availability on biomass allocation patterns in 27 species of herbaceous plants

We investigated allocation to roots, stems and leaves of 27 species of herbaceous clonal plants grown at two nutrient levels. Allocation was analyzed as biomass ratios and also allometrically. As in other studies, the fraction of biomass in stems and, to a lesser extent, in leaves, was usually higher in the high-nutrient treatment than in the low-nutrient treatment, and the fraction of biomass in roots was usually higher under low-nutrient conditions. The relationship between the biomass of plant structures fits the general allometric equation, with an exponent 1 in most of the species. The different biomass ratios under the two nutrient conditions represented points on simple allometric trajectories, indicating that natural selection has resulted in allometric strategies rather than plastic responses to nutrient level. In other words, in most of the species that changed allocation in response to the nutrient treatment, these changes were largely a consequence of plant size. Our data suggest that some allocation patterns that have been interpreted as plastic responses to different resource availabilities may be more parsimoniously explained as allometric strategies.     

Root allocation and foraging precision in heterogeneous soils

Root growth patterns respond to small-scale resource heterogeneity and the presence of roots of neighboring plants, but how a plant integrates its responses to these cues is not well understood. In the presence of neighbors, plants may shift allocation to roots as a consequence of plant size and root:shoot allometry, as a response to resource depletion by neighbors, or through a direct response to neighbor presence. The same response pathways also have the potential to alter proliferation in resource-rich patches in soil.Four species of grassland plants were grown in the greenhouse as single plants, monocultures, and mixtures. Root length allocation as a function of shoot mass was examined for background soil and fertilized patches. Plants grown with same-species neighbors followed the same allometric trajectory as single plants for root length in background soil, so any change in root allocation was due only to reduced plant size. Root proliferation in patches declined with neighbors, consistent with a response to resource depletion. Mixtures overproduced roots in both background soil and in patches, relative to plants of the same size in monocultures.     

The effect of allometric partitioning on herbivory tolerance in four species in South China

Herbivory tolerance can offset the negative effects of herbivory on plants and plays an important role in both immigration and population establishment. Biomass reallocation is an important potential mechanism of herbivory tolerance. To understand how biomass allocation affects plant herbivory tolerance, it is necessary to distinguish the biomass allocations resulting from environmental gradients or plant growth. There is generally a tight balance between the amounts of biomass invested in different organs, which must be analyzed by means of an allometric model. The allometric exponent is not affected by individual growth and can reflect the changes in biomass allocation patterns of different parts. Therefore, the allometric exponent was chosen to study the relationship between biomass allocation pattern and herbivory tolerance. We selected four species (Wedelia chinensis, Wedelia trilobata, Merremia hederacea, and Mikania micrantha), two of which are invasive species and two of which are accompanying native species, and established three herbivory levels (0%, 25% and 50%) to compare differences in allometry. The biomass allocation in stems was negatively correlated with herbivory tolerance, while that in leaves was positively correlated with herbivory tolerance. Furthermore, the stability of the allometric exponent was related to tolerance, indicating that plants with the ability to maintain their biomass allocation patterns are more tolerant than those without this ability, and the tendency to allocate biomass to leaves rather than to stems or roots helps increase this tolerance. The allometric exponent was used to remove the effects of individual development on allocation pattern, allowing the relationship between biomass allocation and herbivory tolerance to be more accurately explored. This research used an allometric model to fit the nonlinear process of biomass partitioning during the growth and development of plants and provides a new understanding of the relationship between biomass allocation and herbivory tolerance.     

Determination of the polymorphic composition of smooth pea starch

A method for estimating the proportions of ‘A’ and ‘B’ polymorphs comprising a sample of ‘C’ type starch is proposed which uses established experimental techniques with commercially available spreadsheet and X-ray analysis software. Waxy maize, potato and smooth pea starches were used to provide X-ray diffraction patterns characteristic of the ‘A’, ‘B’ and ‘C’ starch polymorphs. Samples of amorphous starches were also prepared. The method initially involved subtraction of the amorphous phase and instrumental background from the X-ray diffraction patterns of each starch sample using the spreadsheet program, Lotus 1-2-3. The remainder of the pattern, representing the crystalline portion of the starch sample, was then analysed by profile fitting to elucidate the positions and areas of individual diffraction peaks. The ratio of the total peak area to the areas under peaks characteristic of ‘A’ and ‘B’ type starches, respectively, were used to calculate the relative proportions of these polymorphs in smooth pea starch. These proportions were found to be 56±3% ‘A’ polymorph to 44±3% ‘B’ polymorph. A ‘C’ type pattern was constructed by using Lotus 1-2-3 to combine diffraction patterns from the crystalline portions of ‘A’ and ‘B’ type starches in the proportions given above. Polymorph patterns were obtained by manipulation of the diffraction patterns from the crystalline portions of starches using Lotus 1-2-3. An ‘A’ type pattern was obtained by subtraction of a ‘B’ type pattern from that of a ‘C’ type. Similarly, a ‘B’ type pattern was obtained by subtraction of an ‘A’ type pattern from that of a ‘C’ type.     

Short-term effects of fire and competition on growth and plasticity of the yellow pitcher plant,Sarracenia alata (Sarraceniaceae)

Although recurrent fires are widely assumed to reduce competitive interference of plants of pine savannas, rarely has this assumption been tested explicitly. This 2-yr study reports on the interactive effects of fire and neighbors on short-term growth responses and plasticity in allocation patterns of a carnivorous plant, the yellow pitcher plant, Sarracenia alata. This species relies upon pitfall traps (pitchers) to attract and capture insects to obtain nutrients. Neighbors reduced the growth rate of individual ramet transplants (phytometers) in one but not both years of the study. The effect of neighbors on total (i.e., both above- and belowground) productivity of phytometers was not reduced by a winter fire. Neighbors had a greater effect on large plants than on small plants. Although fire did not affect the growth rate of phytometers in the short term, allocation patterns were greatly altered by both neighbors and fire. Allocation to pitchers increased at the expense of belowground organs following fire and in the absence of neighbors at the unburned site. Results of the current study suggest that adult pitcher plants may tolerate competition from neighboring vegetation by reducing allocation to costly pitchers during years without fire.     

Biomass allocation in old-field annual species grown in elevated CO2 environments: no evidence for optimal partitioning

Increased atmospheric carbon dioxide supply is predicted to alter plant growth and biomass allocation patterns. It is not clear whether changes in biomass allocation reflect optimal partitioning or whether they are a direct effect of increased growth rates. Plasticity in growth and biomass allocation patterns was investigated at two concentrations of CO₂ ([CO₂]) and at limiting and nonlimiting nutrient levels for four fast‐ growing old‐field annual species. Abutilon theophrasti, Amaranthus retroflexus, Chenopodium album, and Polygonum pensylvanicum were grown from seed in controlled growth chamber conditions at current (350 μmol mol^?1, ambient) and future‐ predicted (700 μmol mol^?1, elevated) CO₂ levels. Frequent harvests were used to determine growth and biomass allocation responses of these plants throughout vegetative development. Under nonlimiting nutrient conditions, whole plant growth was increased greatly under elevated [CO₂] for three C3 species and moderately increased for a C4 species (Amaranthus). No significant increases in whole plant growth were observed under limiting nutrient conditions. Plants grown in elevated [CO₂] had lower or unchanged root:shoot ratios, contrary to what would be expected by optimal partitioning theory. These differences disappeared when allometric plots of the same data were analysed, indicating that CO₂‐induced differences in root:shoot allocation were a consequence of accelerated growth and development rates. Allocation to leaf area was unaffected by atmospheric [CO₂] for these species. The general lack of biomass allocation responses to [CO₂] availability is in stark contrast with known responses of these species to light and nutrient gradients. We conclude that biomass allocation responses to elevated atmospheric [CO₂] are not consistent with optimal partitioning predictions.     

Soil temperature drives elevational patterns of reproductive allometry in a biodiversity hotspot

Understanding the geographic patterns of reproductive allocation helps in clarifying the selective forces that shape the reproductive strategies of plants. However, studies on the elevational patterns of reproductive allocation remain limited. Moreover, although soil attributes have long been suspected to drive elevational patterns of reproductive allocation, few studies have explored this relationship. Delaying reproduction and allocating a high proportion of biomass to vegetative organs may be risky for plants living under high-elevation habitats, as these two processes can potentially lead to plant reproductive failure due to the low temperatures and short growing seasons at high elevations. Thus, we hypothesize that reproductive effort will increase with elevation and the elevational pattern of reproductive allocation will be largely driven by soil attributes, given their covariation with elevation. To test these hypotheses, we determined the vegetative and reproductive biomass of individual Impatiens arguta (Balsaminaceae) plants across 12 populations in the Gaoligong Mountains (China), and collected data on soil temperature, nutrients, moisture, and pH for each population. Based on standard major axis regression and linear regression models, we found that (1) both vegetative and reproductive biomass decreased with elevation; (2) all populations demonstrated significant allometric slopes (i.e., linear coefficients of log[reproductive biomass]???log[vegetative biomass] regressions)?>?1; (3) allometric slopes decreased with elevation; and (4) soil temperature was a better predictor of the allometric slope than elevation, i.e., the allometric slope decreased with soil temperature. These results suggest that plant species growing at high elevation invest proportionately more resources to reproduction as an adaptation to low-temperature environments, and reproductive output is heavily dependent on vegetative growth. This study provides the first evidence of soil temperature driving reproductive allocation patterns, which suggests that plant species will favor allocation to growth under increasing soil temperatures with climate warming.

     

不同密度猪毛菜形态结构性状及生物量分配策略的异速关系

植物的生长特性随环境条件的变化具有可塑性,而不同的环境因素对植物可塑性的影响也不尽相同。利用异速分析的方法,通过模拟退化草地恢复过程中猪毛菜（Salsola collina）的不同种群密度（16、44、100、400株/m²）,研究其形态结构性状及生物量分配策略的异速关系在种群密度间的差异。结果表明,种群密度增大能抑制猪毛菜的生长,而且对猪毛菜的株高、根长、一级分枝数、二级分枝数、三级分枝数以及总分枝长均产生了极显著的影响,表明种群密度的变化使得植物的高生长和侧向生长发生了显著变化。种群密度的变化也引起了植物生物量的变化,其中植物根、茎、叶间的生物关系是一种表观可塑性,植物的生长策略未发生改变,只是植物个体大小发生改变引起的生物量分配的变化。植物株高、总分枝长、一级分枝数及繁殖分配的变化,是由种群密度变化引起的,植物的适应策略发生了改变,是真正的可塑性。种群密度改变了植物的繁殖分配策略,而未改变植物叶的分配策略,说明当环境发生变化时,植物调整了其繁殖策略以适应环境因素的改变,以保证种群的生存繁衍。     

Can heterochrony help explain the high morphological diversity within the genus Niphargus (Crustacea: Amphipoda)?

Postembryonic development of nine species of Niphargus (Crustacea: Amphipoda) was studied, with spine development and shifts in allometric growth being combined in developmental sequences that were compared across species. The developmental sequences show high diversity with respect to the position of individual events in the sequence, as well as a high frequency of events being inapplicable in some species. Within Niphargus, the highest degree of independence between events occurs mainly in early mid-aged instars, where the shifts in position are both the largest in magnitude and also the most frequent in occurrence. Constructive troglomorphic features of subterranean species were inferred to develop more because of accelerated growth rather than a delayed offset of growth. Shifts in both relative timing and growth rate appeared to have played a role in the evolution of sexually dimorphic elongated appendages. Growth patterns differed greatly between species for individual sexually dimorphic and troglomorphic traits, hinting at their possible independent origin. The independence between developmental events that was generally apparent might indicate the existence of an important genetic basis for the extreme intrageneric morphological variation arising from numerous highly variable body parts being combined in a ‘mosaic’ manner. Moreover, we suggest that the overall diversity of the genus might actually be underestimated in light of possible convergent features accompanied by ‘cryptic’ speciation. We conclude that both sequential and growth heterochrony appear to have played a key role in the evolution of Niphargus, the most diverse genus of freshwater amphipods. A preliminary list of heterochronic characters is provided.     

Transitions between cooperative and non-cooperative responding in the 'Pigeon's Dilemma'

Two pigeons played Iterated Prisoner's Dilemma (IPD) against a simulated opponent pre-programmed to play ‘Tit-For-Tat’ (TFT) and ‘Random’ (RND) strategies. Each pigeon received differential amounts of access to food following choices of either ‘cooperate’ or ‘defect’ on a trial. After 1000 trials against TFT and 500 trials against RND, results indicated that choice allocation was optimal when the birds played against RND but was sub-optimal when the birds played against TFT. In order to determine why the pigeons responded suboptimally against TFT, a trial-by-trial analysis of the data was conducted. The analysis revealed that once a pigeon had received the ‘Sucker's’ payoff (S), it was more likely to defect and receive the ‘Temptation’ payoff (T) than to cooperate and receive the ‘Reward’ (R) payoff. Local reinforcement contingencies appear to determine suboptimal responding against TFT in the iterated Pigeon's Dilemma.     

Competitive regulation of plant allometry and a generalized model for the plant self-thinning process

Taking into account the individual growth form (allometry) in a plant population and the effects of intraspecific competition on allometry under the population self-thinning condition, and adopting Ogawa's allometric equation 1/y = 1/axb + 1/c as the expression of complex allometry, the generalized model describing the change mode of r (the self-thinning exponential in the self-thinning equation, log M = K + log N, where M is mean plant mass, K is constant, and N is population density) was constructed. Meanwhile, with reference to the changing process of population density to survival curve type B, the exponential, r, was calculated using the software MATHEMATICA 4.0. The results of the numerical simulation show that (1) the value of the self-thinning exponential, r, is mainly determined by allometric parameters; it is most sensitive to change of b of the three allometric parameters, and a and c take second place; (2) the exponential, r, changes continuously from about -3 to the asymptote -1; the slope of -3/2 is a transient value in the population self-thinning process; (3) it is not a 'law' that the slope of the self-thinning trajectory equals or approaches -3/2, and the long-running dispute in ecological research over whether or not the exponential, r, equals -3/2 is meaningless. So future studies on the plant self-thinning process should focus on investigating how plant neighbor competition affects the phenotypic plasticity of plant individuals, what the relationship between the allometry mode and the self-thinning trajectory of plant population is and, in the light of evolution, how plants have adapted to competition pressure by plastic individual growth.     

Orthology,paralogy and proposed classification for paralog subtypes

The paper clears up confusions about the concepts of orthology and paralogy, particularly in cases involving gene family expansions. The terms ‘inparalog’ and ‘outparalog’ are defined to distinguish ancient paralogs from lineage-specific ones.     

Species traits and shoot-root biomass allocation in 20 dry-grassland species

Aims A plant has a limited amount of resources at any time and it allocates them to different structures. In spite of the large number of previous studies on allocation patterns within single species, knowledge of general patterns in species allocation is still very limited. This is because each study was done in different conditions using different methodology, making generalization difficult. We investigate intraspecific above- versus below-ground biomass allocation among individuals across a spectrum of dry-grassland plant species at two different developmental stages and ask whether allocation is age- and species specific, and whether differences among species can be explained by their life-history traits and phylogeny.Methods We collected data on above- and below-ground biomass of seedlings and adult plants of 20 species from a common garden experiment. We analysed data on shoot–root biomass allocation allometrically and studied the relationship between the allometric exponents (slopes on log–log scale), species life-history traits and phylogenetic distances.Important findings We found isometric as well as allometric patterns of biomass allocation in the studied species. Seedlings and adult individuals of more than half of the species differed in their above- versus below-ground biomass allometric exponents. Seedlings and adult individuals of the remaining species differed in their allometric coefficients (intercepts). Annual species generally allocated proportionally more to above- than below-ground biomass as seedlings than as adults, whereas perennial species showed the opposite pattern. Plant life-history traits, such as plant life span, age of first flowering, month in which the species begin flowering and specific leaf area were much more important in explaining differences in shoot–root allometry among species than were phylogenetic relationships. This suggests that allocation patterns vary greatly among closely related species but can be predicted based on species life-history traits.     

Biosynthesis of Hb in individual fetal liver bursts : γ-Chain production peaks earlier than β-chain in the erythropoietic pathway

The relative synthesis of α-, β-, ^Gγ- and ^Aγ-globin chains has been evaluated in single fetal liver bursts, which were grown in methylcellulose cultures, individually labelled with [³H]leucine and then analysed via iso-electric focusing. Well-hemoglobinized bursts demonstrate a homogeneous globin synthetic pattern, characterized by prevalent HbF (+some HbA) synthesis: thus, they apparently originate from a homogeneously programmed population of erythroid burst-forming unit (BFU-E). On day 8–9 of culture, the synthetic pattern in ‘mature’ (i.e., well-hemoglobinized) bursts has been compared with that in simultaneously-grown, ‘immature’ (i.e., poorly-hemoglobinized) colonies. These patterns have been further compared with that in ‘matured’ bursts (identified in situ as immature on day 8–9 and labelled 2–4 days later when matured). The ‘immature’ colonies showed very low levels of relative β-globin synthesis, while the ‘mature’ ones demonstrated a more elevated production of β-chain. Significantly, the ‘matured’ bursts showed a globin chain synthetic pattern similar to that of previously labelled ‘matured’ colonies. It is postulated therefore that in fetal liver (and also in adult marrow) the synthesis of γ-chain is linked to an early differentiation stage of erythroblasts, while β-globin synthesis is largely activated at a more advanced maturation stage.     

Size inequality and the tragedy of the commons phenomenon in plant competition

Game theory predicts that the evolutionarily stable level of root production is greater for plants grown with neighbours compared to plants grown alone, even when the available resources per plant are constant. This follows from the fact that for plants grown alone, new roots compete only with other roots on the same plant, whereas for multiple plants grown in a group, new roots can also compete with the roots of other plants, thereby potentially acquiring otherwise unavailable resources at their neighbours’ expense. This phenomenon, which results in plants grown with neighbours over-proliferating roots at the expense of above-ground biomass, has been described as a ‘tragedy of the commons’, and requires that plants can distinguish self from non-self tissues. While this game theoretical model predicts the evolutionarily stable strategies of individual plants, it has only been tested on average allocation patterns of groups of plants. This is problematic, because average patterns can appear to reflect a tragedy of the commons, even when none has occurred. In particular, assuming (1) a decelerating relationship between individual plant biomass and the amount of resources available, and (2) greater size inequality in plants grown with neighbours compared to plants grown alone (due to asymmetric competition), then plants grown with neighbours should, at least on average, be smaller than plants grown alone. This is a manifestation of ‘Jensen’s Inequality’, which states that for decelerating functions, the average value of the function is less than the function of the average value. We suggest that Jensen’s Inequality should serve as an appropriate null hypothesis for examining biologically-based explanations of changes in biomass allocation strategies.