ABSTRACT
Most cells tightly control the length of their cilia. The regulation likely involves intraflagellar transport (IFT), a bidirectional motility of multi-subunit particles organized into trains that deliver building blocks into the organelle. In Chlamydomonas, the anterograde IFT motor kinesin-2 consists of the motor subunits FLA8 and FLA10 and the nonmotor subunit KAP. KAP dissociates from IFT at the ciliary tip and diffuses back to the cell body. This observation led to the diffusion-as-a-ruler model of ciliary length control, which postulates that KAP is progressively sequestered into elongating cilia because its return to the cell body will require increasingly more time, limiting motor availability at the ciliary base, train assembly, building block supply, and ciliary growth. Here, we show that Chlamydomonas FLA8 also returns to the cell body by diffusion. However, more than 95% of KAP and FLA8 are present in the cell body and, at a given time, just ~1% of the motor participates in IFT. After repeated photobleaching of both cilia, IFT of fluorescent kinesin subunits continued indicating that kinesin-2 cycles from the large cell-body pool through the cilia and back. Furthermore, growing and full-length cilia contained similar amounts of kinesin-2 subunits and the size of the motor pool at the base changed only slightly with ciliary length. These observations are incompatible with the diffusion-as-a-ruler model, but rather support an "on-demand model," in which the cargo load of the trains is regulated to assemble cilia of the desired length.
ABSTRACT
Black carbon (BC), an important component of organic carbon (OC) produced from incomplete combustion of carbon compounds, is widespread and affects the global carbon storage. The objectives of this study were to analyze the BC contents and fluxes in the last 150 years to determine the causes of differences in the three profiles of the Songnen Plain of Northeast China and to estimate the BC storage in the wetlands of the Songnen Plain. In the three sampling sites, BC fluxes in the period between 1950 and the present time increased by the ratios of 1.3, 31.1 and 1.4, respectively, compared to their own baseline between 1850 and 1900. Furthermore, the BC fluxes varying from 0.76 to 5.63 g m-2 y-1 in the three profiles had an opposite trend with the sand percentages with mean values changing from 78.9% to 19.6%, suggesting that sand desertification might additionally affect the BC processes in the region.
