The biomechanics of pollen release: new perspectives on the evolution of wind pollination in angiosperms

Evolutionary transitions from animal to wind pollination have occurred repeatedly during the history of the angiosperms, but the selective mechanisms remain elusive. Here, we propose that knowledge of pollen release biomechanics is critical for understanding the ecological and evolutionary processes underpinning this shift in pollination mode. Pollen release is the critical first stage of wind pollination (anemophily) and stamen properties are therefore likely to be under strong selection early in the transition. We describe current understanding of pollen release biomechanics to provide insights on the phenotypic and ecological drivers of wind pollination. Pollen release occurs when detachment forces dominate resistive forces retaining pollen within anthers. Detachment forces can be active or passive depending on whether they require energy input from the environment. Passive release is more widespread in anemophilous species and involves processes driven by steady or unsteady aerodynamic forces or turbulence-induced vibrations that shake pollen from anthers. We review empirical and theoretical studies suggesting that stamen vibration is likely to be a key mechanism of pollen release. The vibration response is governed by morphological and biomechanical properties of stamens, which may undergo divergent selection in the presence or absence of pollinators. Resistive forces have rarely been investigated for pollen within anthers, but are probably sensitive to environmental conditions and depend on flower age, varying systematically between animal- and wind-pollinated species. Animal and wind pollination are traditionally viewed as dichotomous alternatives because they are usually associated with strikingly different pollination syndromes. But this perspective has diverted attention from subtler, continuously varying traits which mediate the fluid dynamic process of pollen release. Reinterpreting the flower as a biomechanical entity that responds to fluctuating environmental forces may provide a promising way forward. We conclude by identifying several profitable areas for future research to obtain deeper insight into the evolution of wind pollination.     

Seed size,pollination costs and angiosperm success

Summary Seed plants capture pollen before seeds are dispersed and abort unpollinated ovules. As a result, each seed is associated with an accessory cost that represents the costs of pollen capture and the costs of aborted ovules. Accessory costs may explain the minimum seed size among species, because these costs are likely to comprise a greater proportion of total reproductive allocation in species with smaller seeds. For very small propagules, the costs of pollination may not be worth the benefits, perhaps explaining the persistence of pteridophytic reproduction at small propagule sizes. The smallest angiosperm seeds are much smaller than the smallest gymnosperm seeds, both in the fossil record and in the modern flora. This suggests that angiosperms can produce pollinated ovules more cheaply than gymnosperms. Pollination becomes less efficient as a species decreases in abundance, and this loss of efficiency is greater for species with a higher accessory cost per seed. We propose that the greater reproductive efficiency of angiosperms when rare can explain why angiosperm-dominated floras were more speciose than the gymnosperm-dominated floras they replaced.     

沙丘稀有种准噶尔无叶豆花部综合特征与传粉适应性

植物花部特征进化与传粉适应性一直是进化生态学领域关注的核心问题之一。以古尔班通古特沙漠自然生长的准噶尔无叶豆为对象,对其花部特征和传粉特性进行了野外观察和室内的分析研究。结果表明:种群花期历时21 d,花序花期历时7-12 d,单花花期一般3 d,若遇阴雨天气,花期可延长1-2 d,整个花期龙骨瓣一直保持闭合状态。单花10枚花药在旗瓣微张时已全部完成散粉。准噶尔无叶豆主要靠分泌花蜜、鲜艳的花色以及旗瓣基部的黄色辐射状纹理结构吸引传粉者。准噶尔无叶豆花期的有效传粉者为4种蜂类昆虫,它们的平均访花频率为(7.75±0.57)次·花^-1·d^-1,访花高峰期表现为三峰型: 13:00-14:00,16:00-17:00和19:00-20:00。准噶尔无叶豆人工套袋实验表明该种为自交亲和型,主动自交少见,生殖成功依赖传粉者。胚珠成功受精至果实完全成熟阶段存在自交衰退,柱头角质层结构和花粉刷结构是准噶尔无叶豆在进化过程中形成的减少自交,倾向异交的机制。     

Phylogenetics of tribe Orchideae (Orchidaceae: Orchidoideae) based on combined DNA matrices: inferences regarding timing of diversification and evolution of pollination syndromes

Background and aims

Tribe Orchideae (Orchidaceae: Orchidoideae) comprises around 62 mostly terrestrial genera, which are well represented in the Northern Temperate Zone and less frequently in tropical areas of both the Old and New Worlds. Phylogenetic relationships within this tribe have been studied previously using only nuclear ribosomal DNA (nuclear ribosomal internal transcribed spacer, nrITS). However, different parts of the phylogenetic tree in these analyses were weakly supported, and integrating information from different plant genomes is clearly necessary in orchids, where reticulate evolution events are putatively common. The aims of this study were to: (1) obtain a well-supported and dated phylogenetic hypothesis for tribe Orchideae, (ii) assess appropriateness of recent nomenclatural changes in this tribe in the last decade, (3) detect possible examples of reticulate evolution and (4) analyse in a temporal context evolutionary trends for subtribe Orchidinae with special emphasis on pollination systems.

Methods

The analyses included 118 samples, belonging to 103 species and 25 genera, for three DNA regions (nrITS, mitochondrial cox1 intron and plastid rpl16 intron). Bayesian and maximum-parsimony methods were used to construct a well-supported and dated tree. Evolutionary trends in the subtribe were analysed using Bayesian and maximum-likelihood methods of character evolution.

Key Results

The dated phylogenetic tree strongly supported the recently recircumscribed generic concepts of Bateman and collaborators. Moreover, it was found that Orchidinae have diversified in the Mediterranean basin during the last 15 million years, and one potential example of reticulate evolution in the subtribe was identified. In Orchidinae, pollination systems have shifted on numerous occasions during the last 23 million years.

Conclusions

The results indicate that ancestral Orchidinae were hymenopteran-pollinated, food-deceptive plants and that these traits have been dominant throughout the evolutionary history of the subtribe in the Mediterranean. Evidence was also obtained that the onset of sexual deception might be linked to an increase in labellum size, and the possibility is discussed that diversification in Orchidinae developed in parallel with diversification of bees and wasps from the Miocene onwards.     

Functional homogenization of flower visitor communities with urbanization

Land‐use intensification and resulting habitat loss are put forward as the main causes of flower visitor decline. However, the impact of urbanization, the prime driver of land‐use intensification in Europe, is poorly studied. In particular, our understanding of whether and how it affects the composition and functioning of flower visitor assemblages is scant, yet required to cope with increasing urbanization worldwide. Here, we use a nation‐wide dataset of plant‐flower visitor (Coleoptera, Diptera, Hymenoptera, Lepidoptera) interactions sampled by citizen scientists following a standardized protocol to assess macroecological changes in richness and composition of flower visitor communities with urbanization. We measured the community composition by quantifying the relative occurrence of generalist and specialist flower visitors based on their specialisation on flowering plant families. We show that urbanization is associated with reduced flower visitor richness and a shift in community composition toward generalist insects, indicating a modification of the functional composition of communities. These results suggest that urbanization affects not only the richness of flower visitor assemblages but may also cause their large‐scale functional homogenization. Future research should focus on designing measures to reconcile urban development with flower visitor conservation.     

Optimization of compressive loading parameters to mimic in vivo cervical spine kinematics in vitro

The human cervical spine supports substantial compressive load in vivo. However, the traditional in vitro testing methods rarely include compressive loads, especially in investigations of multi-segment cervical spine constructs. Previously, a systematic comparison was performed between the standard pure moment with no compressive loading and published compressive loading techniques (follower load – FL, axial load – AL, and combined load – CL). The systematic comparison was structured a priori using a statistical design of experiments and the desirability function approach, which was chosen based on the goal of determining the optimal compressive loading parameters necessary to mimic the segmental contribution patterns exhibited in vivo. The optimized set of compressive loading parameters resulted in in vitro segmental rotations that were within one standard deviation and 10% of average percent error of the in vivo mean throughout the entire motion path. As hypothesized, the values for the optimized independent variables of FL and AL varied dynamically throughout the motion path. FL was not necessary at the extremes of the flexion–extension (FE) motion path but peaked through the neutral position, whereas, a large negative value of AL was necessary in extension and increased linearly to a large positive value in flexion. Although further validation is required, the long-term goal is to develop a “physiologic” in vitro testing method, which will be valuable for evaluating adjacent segment effect following spinal fusion surgery, disc arthroplasty instrumentation testing and design, as well as mechanobiology experiments where correct kinematics and arthrokinematics are critical.     

Pollination and reproductive system of synchronopatric species of Cactaceae (Cactoideae) subject to interspecific flow of pollen: an example of ecological adaptation in the Brazilian Chaco

• Three synchronopatric Cactaceae species, Echinopsis rhodotricha, Harrisia balansae and Praecereus saxicola, have mostly nocturnal anthesis and similar flowers, characteristics that motivated us to perform a comparative study of reproductive ecology.
• Reproductive phenology was sampled monthly from December 2014 to November 2015. We describe floral biology, breeding system via pollination treatments and evaluate floral visitors from focal and filming observations. Pollen grains found on moth proboscis were compared among cactus species under light microscopy. We used fluorescent dye particles to test intra‐ and interspecific pollen flow.
• These three species have extended flowering with greater intensity in the wet season, causing high overlap. They have white and hypocrateriformis flowers that open at twilight or nightfall and last about 15 h. H. balansae seems to be self‐incompatible, while E. rhodotricha presented self‐compatibility. P. saxicola presented self‐fertility, but most of the population seems to be self‐incompatible. We suggest sphingophily for the three species, but only P. saxicola was visited by Manduca rustica (Sphingidae). However, we observed pollen grains of all three species on the proboscis of moths, especially M. rustica and M. sexta. Prolonged anthesis allowed bees (herein considered as secondary pollinators) to visit flowers of E. rhodotricha and P. saxicola.
• It can be concluded that the studied species share nocturnal and diurnal pollinators, suggesting interspecific pollen flow, which, however, could not be detected with fluorescent dye particles. In view of the low frequency of primary pollinators, it appears that these three species have different reproductive strategies, ensuring the fruiting and production of viable seeds.

     

The 141-year period for Dr. Beal's seed viability experiment: A hybrid surprise

Premise

In 1879, Dr. William Beal buried 20 glass bottles filled with seeds and sand at a single site at Michigan State University. The goal of the experiment was to understand seed longevity in the soil, a topic of general importance in ecology, restoration, conservation, and agriculture, by periodically assaying germinability of these seeds over 100 years. The interval between germination assays has been extended and the experiment will now end after 221 years, in 2100.

Methods

We dug up the 16th bottle in April 2021 and attempted to germinate the 141-year-old seeds it contained. We grew germinants to maturity and identified these to species by vegetative and reproductive phenotypes. For the first time in the history of this experiment, genomic DNA was sequenced to confirm species identities.

Results

Twenty seeds germinated over the 244-day assay. Eight germinated in the first 11 days. All 20 belonged to the Verbascum genus: Nineteen were V. blattaria according to phenotype and ITS2 genotype; and one had a hybrid V. blattaria × V. thapsus phenotype and ITS2 genotype. In total, 20/50 (40%) of the original Verbascum seeds in the bottle germinated in year 141.

Conclusions

While most species in the Beal experiment lost all seed viability in the first 60 years, a high percentage of Verbascum seeds can still germinate after 141 years in the soil. Long-term experiments such as this one are rare and invaluable for studying seed viability in natural soil conditions.     

河南芝麻授粉昆虫初步调查

为了清楚了解芝麻花授粉昆虫种类、昆虫的授粉规律和授粉时间,2009年对河南省平舆县高杨店乡的13个样点芝麻花授粉昆虫进行调查,采集到授粉昆虫标本65份.经过整理鉴定,芝麻花授粉昆虫共计5目14科25种.其中,蜜蜂占授粉昆虫的92%,是大田芝麻主要的授粉昆虫.蜜蜂授粉出勤高峰出现在8:00～8:50和18:00～18:50.